Restoring Western Ranges and Wildlands

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United States Department of Agriculture Restoring Western Forest Service Rocky Mountain Research Station General Technical Ranges and Wildlands Report RMRS-GTR-136-vol. 2 September 2004 Volume 2 Chapters 18-23, Index Abstract ______

Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps. 2004. Restoring western ranges and wildlands. Gen. Tech. Rep. RMRS-GTR-136-vol-2. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. Pages 295Ð 698 plus index.

This work, in three volumes, provides background on philosophy, processes, plant materials selection, site preparation, and seed and seeding equipment for revegetating disturbed rangelands, emphasizing use of native species. The 29 chapters include guidelines for planning, conducting, and managing, and contain a compilation of rangeland revegetation research conducted over the last several decades to aid practitioners in reestablishing healthy communities and curbing the spread of invasive species. Volume 2 contains chapters 18-23 plus the index.

Keywords: rehabilitation, revegetation, plant ecology, seed, plant communities, wildlife habitat, invasive species, equipment, plant materials, native plants

A B

A—Curlleaf mountain mahogany. B—Arrowleaf balsamroot. C—Bluebunch wheatgrass seed production field. D—Antelope bitterbrush achene. E—Squirreltail. Restoring Western Ranges and Wildlands Compilers Stephen B. Monsen Volume 2 Richard Stevens Nancy L. Shaw Chapters 18–23, index

D E

i The Compilers ______Stephen B. Monsen (retired), Botanist, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah Richard Stevens, Project Leader (retired), Utah Division of Wildlife Resources, Great Basin Research Center, Ephraim, Utah Nancy L. Shaw, Research Botanist, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Boise, Idaho

The Authors ______James N. Davis is Project Leader, Big Game Range Trend Program, Utah Division of Wildlife Resources, Shrub Sciences Laboratory, Provo, Utah. Sherel K. Goodrich is Ecologist, U.S. Department of Agriculture, Forest Service, Ashley National Forest, Vernal, Utah. Kent R. Jorgensen was Wildlife Biologist (deceased), Great Basin Research Center, Utah Division of Wildlife Resources, Ephraim, Utah. Carlos F. Lopez was Soil Scientist, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. E. Durant McArthur is Project Leader and Research Geneticist, U.S. Department of Agricul- ture, Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. Stephen B. Monsen was Botanist (retired), U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. He is currently a private natural resources consultant and volunteer worker for the Rocky Mountain Research Station. David L. Nelson was Plant Pathologist (retired), U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. He continues contractual and volunteer work for the Rocky Mountain Research Station.

Nancy L. Shaw is Research Botanist, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Boise, Idaho. Richard Stevens was Project Leader (retired), Great Basin Research Center, Utah Division of Wildlife Resources, Ephraim, Utah. Arthur R. Tiedemann was Project Leader and Range Scientist (retired), U.S. Department of Agriculture, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. John F. Vallentine was Professor (Emeritus), Department of Integrative Biology, Brigham Young University, Provo, Utah. Gordon A. Van Epps was Associate Professor (retired), Snow Field Station, Utah Agricultural Experiment Station, Utah State University, Ephraim, Utah. Bruce L. Welch is Plant Physiologist, U.S. Department of Agriculture, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. Steven G. Whisenant is Professor, Department of Rangeland Ecology and Management, Texas A&M University, College Station, Texas. G. Richard Wilson was Director (retired), Utah State Seed Laboratory, Utah Department of Agriculture, Salt Lake City, Utah.

ii Contents

Volume 1: Page Foreword by E. Durant McArthur ...... iii 1. History of Range and Wildlife Habitat Restoration in the Intermountain West ...... 1 2. The Intermountain Setting ...... 7 3. Research Background ...... 15 4. Basic Considerations for Range and Wildland Revegetation and Restoration...... 19 5. Restoration or Rehabilitation Through Management or Artificial Treatments ...... 25 6. Climate and Terrain ...... 33 7. Assessing Soil Factors in Wildland Improvement Programs ...... 39 8. Controlling Plant Competition ...... 57 9. Mechanical Plant Control ...... 65 10. Herbicides for Plant Control ...... 89 11. Vegetative Manipulation with Prescribed Burning ...... 101 12. Seedbed Preparation and Seeding Practices ...... 121 13. Incorporating Wildlife Habitat Needs into Restoration and Rehabilitation Projects ...... 155 14. Nutritive Principles in Restoration and Management ...... 175 15. Plant Pathology and Managing Wildland Plant Disease Systems ...... 187 16. Management of Restored and Revegetated Sites ...... 193 17. Guidelines for Restoration and Rehabilitation of Principal Plant Communities ...... 199

Volume 2: 18. Grasses ...... 295 19. Forbs for Seeding Range and Wildlife Habitats ...... 425 20. Chenopod Shrubs ...... 467 21. Composite Shrubs ...... 493 22. Rosaceous Shrubs ...... 539 23. Shrubs of Other Families ...... 597

Volume 3: 24. Seed Collection, Cleaning, and Storage ...... 699 25. Shrub and Forb Seed Production ...... 717 26. Seed Germination ...... 723 27. Seed Testing Requirements and Regulatory Laws ...... 733 28. Establishing Plants by Transplanting and Interseeding ...... 739 29. Production and Use of Planting Stock ...... 745 References ...... 769 Appendix 1 Scientific/Common Names ...... 847 Appendix 2 Common/Scientific Names ...... 866 Index ...... Index-1

iii The use of trade or firm names in this publication is for reader information and does not imply endorsement by the U.S. Department of Agriculture of any product or service

Pesticide Precautionary Statement

This publication reports research involving pesticides. It does not contain recommendations for their use, nor does it imply that the uses discussed here have been registered. All uses of pesticides must be registered by appropriate State and/or Federal agencies before they can be recommended.

CAUTION: Pesticides can be injurious to humans, domestic animals, desirable plants, and fish or other wildlife—if they are not handled or applied properly. Use all pesticides selectively and carefully. Follow recommended practices for the disposal of surplus pesticides and pesticide containers.

CAUTION: PESTICIDES

You may order additional copies of this publication by sending your mailing information in label form through one of the following media. Please specify the publication title and number.

Telephone (970) 498-1392 FAX (970) 498-1396 E-mail [email protected] Web site http://www.fs.fed.us/rm Mailing Address Publications Distribution Rocky Mountain Research Station 240 West Prospect Road Fort Collins, CO 80526 Stephen B. Monsen Richard Stevens Chapter Nancy Shaw 18

Grasses

Grasses are adapted to a wide range of edaphic and climatic conditions and are found in nearly all plant communities. In the Western United States, grasses are seeded on disturbances to provide forage (Hull and Holmgren 1964; Vallentine 1989), wildlife habitat (Plummer and others 1968), and watershed stability (Cornelius 1946; Hafenrichter and others 1949; Piper 1934; Stewart and Young 1939). A number of introduced grass species proved well-suited to Western rangelands and received extensive use in early reseeding efforts (Barnes and others 1995; Hafenrichter and others 1968; Moser and others 1996). Use of these and other introductions continued over time, but by the late 1900s, greater emphasis was being placed on the use of native grasses (Roundy and others 1997). Selection of introduced and native grasses for Western rangelands began in the late nineteenth century. Excessive livestock grazing across the West followed early European settlement, resulting in widespread disturbances, weed invasions, and a need for revegetation species to stabilize watersheds and produce forage. A few species and accessions were introduced in the West before 1900 (Hanson 1959). Some arrived by chance (Weintraub

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 295 Chapter 18 Grasses

Chapter Contents by Species

Agropyron cristatum (Fairway Crested Wheatgrass) ...... 297 Agropyron dasystachyum (Thickspike Wheatgrass) ...... 318 Agropyron desertorum (Standard Crested Wheatgrass or Desert Crested Wheatgrass) ...... 320 Agropyron elongatum (Tall Wheatgrass) ...... 325 Agropyron intermedium (Intermediate Wheatgrass) ...... 327 Agropyron sibiricum (Siberian Wheatgrass) ...... 332 Agropyron smithii (Western Wheatgrass or Bluestem Wheatgrass) ...... 334 Agropyron spicatum (Bluebunch Wheatgrass)...... 337 Agropyron trachycaulum (Slender Wheatgrass) ...... 342 Agrostis stolonifera (Redtop or Carpet Bentgrass) ...... 345 Alopecurus arundinaceus (Creeping Foxtail or Reed Foxtail) ...... 346 Alopecurus pratensis (Meadow Foxtail) ...... 346 Aristida purpurea (Red Threeawn, Purple Threeawn) ...... 349 Arrhenatherum elatius (Tall Oatgrass) ...... 351 Bouteloua curtipendula (Sideoats Grama) ...... 353 Bouteloua eriopoda (Black Grama) ...... 355 Bouteloua gracilis (Blue Grama) ...... 356 Bromus anomalus (Nodding Brome) ...... 358 Bromus ciliatus (Fringed Brome) ...... 358 Bromus carinatus (Mountain Brome, California Brome) ...... 360 Bromus inermis (Smooth Brome, Hungarian Brome) ...... 363 Bromus riparius (Meadow Brome) ...... 365 Dactylis glomerata (Orchardgrass) ...... 367 Deschampsia caespitosa (Tufted Hairgrass) ...... 369 Distichlis spicata (Inland Saltgrass) ...... 371 Elymus canadensis (Canada Wildrye) ...... 373 Elymus cinereus (Great Basin Wildrye) ...... 374 Elymus glaucus (Blue Wildrye) ...... 377 Elymus junceus (Russian Wildrye) ...... 379 Elymus salinus (Salina Wildrye) ...... 381 Elymus simplex (Alkali Wildrye or Low-Creeping Wildrye) ...... 382 Elymus triticoides (Creeping Wildrye or Beardless Wildrye) ...... 383 Festuca arundinacea (Tall Fescue, Reed Fescue, or Alta Fescue) ...... 384 Festuca idahoensis (Idaho Fescue or Bluebunch Fescue)...... 386 Festuca ovina (Sheep Fescue or Alpine Fescue)...... 388 Hilaria jamesii (Galleta or Curly Grass)...... 390 Koeleria macrantha (Prairie Junegrass) ...... 392 Oryzopsis hymenoides (Indian Ricegrass) ...... 393 Phalaris arundinacea (Reed Canarygrass) ...... 396 Phleum alpinum (Alpine Timothy or Mountain Timothy) ...... 398 Phleum pratense (Timothy) ...... 399 Poa ampla (Big Bluegrass)...... 401 Poa compressa (Canada Bluegrass) ...... 402 Poa fendleriana (Mutton Bluegrass) ...... 404 Poa pratensis (Kentucky Bluegrass) ...... 406 Poa secunda (Sandberg Bluegrass)...... 408 Sitanion hystrix (Squirreltail, bottlebrush) ...... 410 Sporobolus airoides (Alkali Sacaton) ...... 413 Sporobolus cryptandrus (Gray Sand Dropseed) ...... 414 Stipa columbiana (Subalpine Needlegrass or Columbia Needlegrass) ...... 416 Stipa comata (Needle-and-thread) ...... 418 Stipa lettermanii (Letterman Needlegrass) ...... 419 Stipa thurberiana (Thurber Needlegrass)...... 421 Stipa viridula (Green Needlegrass) ...... 422

296 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

1953); others were selected for specific uses and areas productive species and accessions that might, none- (Dillman 1946). Field testing and seeding trials for theless, be adapted to specific situations. With in- high-elevation disturbances were initiated at this time. creased efforts to reestablish native plant communi- Based on results of these trials, Williams (1898) de- ties, it became essential to more fully utilize the array scribed the values and uses of introduced forage grasses of species and populations adapted to the wide array of for the Eastern Rocky Mountains. environments encountered on Western rangelands. Seeding trials conducted by the Bureau of Plant Many native species have not been brought into seed Industry were initiated in 1902 (Cotton 1908). Forest production, but should be seeded in appropriate areas Service reseeding research utilizing forage species and maintained through proper management. Re- began in 1907 in cooperation with this agency (Sampson search is being conducted to develop seed transfer 1913). Ecological studies of grasses native to the Na- guidelines for native grasses of the Intermountain tional Forests were initiated in about 1911 (USDA region to permit increase of adapted seed supplies for Forest Service 1914). Efforts to evaluate grasses for revegetation that will conserve the genetic diversity of range and wildlife uses were expanded by Forest native populations. Thus, at the present and into the Service researchers and their cooperators by about future, greater consideration will be given to native 1930 (Stewart and others 1939). Cool-season grasses grasses, forbs, and shrubs. received special attention for use on Western range- Grass taxonomy has evolved continuously since lands (Koonce 1946; Piper 1934). Linnaeus published the first descriptions in 1753. Breeding and selection programs were organized Extensive revisions, particularly within the Triticeae, within the USDA Soil Conservation Service, now the have been made over the last few decades as our USDA Natural Resources Conservation Service, in understanding of origins and genetic relationships the 1930s (Hafenrichter and others 1968). The need to increased (Arnow 1987; Barkworth and others 1983). stabilize abandoned farmlands during the 1930’s fur- Löve (1982, 1984) reorganized genera within the ther stimulated the demand for conservation species Triticeae based on genomic relationships. Dewey (1982, (Dillman 1946). Weintraub (1953) published a review 1983a, 1984) concurred with this revision as it pro- of publications on testing and field planting studies vided a clearer picture of phylogenetic and biological conducted between 1920 and 1950. closeness. New taxonomic keys, however, continue to The Soil Conservation Service (now the USDA utilize conventional morphological features to sepa- Natural Resources Conservation Service) and their rate taxa. We subscribe to the recent changes, but we cooperators began releasing grass varieties in the mid have elected to use the more traditional nomenclature 1940s (Alderson and Sharp 1994; Hanson 1959). Some because most land managers, seed dealers, and grow- introduced range and pasture varieties obtained im- ers are better acquainted with it. Traditional and mediate success. Introductions from areas of Europe current nomenclatures are presented in table 3. and Asia with environmental conditions similar to those encountered in Western North America proved useful over wide areas. Testing of drought-tolerant Agropyron cristatum grasses adapted to arid and semiarid rangelands re- Fairway Crested Wheatgrass ______ceived increasing emphasis during recent decades (Barnes and others 1952; Bridges 1942; Christ 1934; Description Forsling and Dayton 1931; Koonce 1946; Piper 1934; Fairway crested wheatgrass, also called crested Price 1938; Stoddart 1946a). Some of the most impor- wheatgrass, is an introduced, long-lived, cool-season, tant grass varieties developed for use on Western range, generally nonrhizomatous bunchgrass that produces wildland, and conservation plantings are listed in table 1. strong tillers and an extensive root system. Stems and Following is a discussion of introduced and native leaves are tightly clustered at the base. Plants are grasses that have proven most useful for seeding generally uniform in stature and growth form. Stem Western wildlands. The range of adaptability, uses, heights within individual plants range from 8 to 20 and characteristics of important grasses are summa- inches (20 to 50 cm), but may reach 40 inches (1 m). rized in table 2. In the past, productive accessions that Stems are generally erect, but some may be bent were palatable and responded well to grazing were slightly at the lower nodes. Leaves develop from the emphasized in plant selection programs for range- plant base and stems. Leaf blades are 2 to 8 inches (5 lands. Characteristics such as good seed production, to 20 cm) long and 0.08 to 0.2 inch (2 to 5 mm) wide, ease of seed cleaning and seeding, and good seedling appearing flat or loosely bent along the midrib. establishment also received priority in the selection Blades are rough, heavily grooved with raised nerves, process (Frischknecht 1951; Hanson and Carnahan and often have a bluish cast. Hairs are sometimes 1956; Plummer and others 1943). Although impor- present on the upper portion of the leaf blade. Auricles tant, these emphases often discouraged the use of less are small and clasping. Ligules are membranous

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 297 Chapter 18 Grasses

(con.)

Attributes

Highly rhizomatous, fine leaves

Early greenup, highly productive

Low growing, nonrhizomatous, winter hardy, selected for

Good forage and seed production, useful on low-fertility

sites of moderate acidity and at high elevations

Dense, low-maintenance turf for erosion control on

Vigorous, highly productive, broad climatic tolerance

Good drought tolerance and provides early greenup,

competitive with annual weeds Rapid development, early maturing, selfseeding, cover crop Tall, productive, used in native seed mixes for wildlife habitat and reestablishment of native plant communities

Erect, vigorous, winter hardy

Excellent establishment and early development, highly Rapid development, erect, leafy, late maturing, heavy

seed and forage producer

Good stand longevity and head smut resistance

NRCS, Pullman, WA Post Falls, ID

Post Falls, ID infertile sites

NRCS, Pullman, WA

NRCS, Lockeford, CA

NRCS, Palmer, AK

NRCS, Aberdeen, ID NRCS, Pullman, WA

Illinois collections

Switzerland Alaska

Canada

Turkey productive, tolerant of shade, winter hardy

Streaker 1982 Combined Jacklin Seed Company,

Gruening 1986 La Cure, NRCS, Palmer, AK

Foothills 2001 Composite of NRCS, Bridger, MT

Germplasm eight accessions

Reubens 1976 Reubens, ID Jacklin Seed Company,

Newport 1958 Lincoln County, OR

Canbar 1979 Blue Mountains, WA

Cucamonga 1949 Cucamonga, CA High Plains 2000 Composite from NRCS, Bridger, MT Germplasm Montana and Wyoming

Service 1989 Whitehorse, Yukon,

Bromar 1946 Whitman County, WA

Garnet 2000 Garnet, MT UCEPC, Meeker, CO

Other cultivars are Fleet and Paddock.

germplasm date Origin seed maintenance

)

—Selected grass cultivars and germplasms for range and wildlife projects.

P. canbyi

Table 1

Common and Cultivar or Release Breeder Bentgrass, redtop scientific names

Bluegrass, big Sherman 1945 Sherman County, OR

Bluegrass, alpine

Bluegrass, Canada

Bluegrass, Kentucky

Bluegrass, Sandberg

( Brome, California Agrostis stolonifera Poa alpina Poa ampla Poa compressa Poa pratensis Poa secunda Bromus carinatus Brome, meadowBromus riparius Brome, mountain Bromus marginatus Regar 1966 Zek, Kars Province,

298 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

(con.)

Attributes

Most widely planted northern type in the West, moderately

spreading type, high yields, good regrowth

High seed yields, excellent seed production

Leafy, persistent, moderate forage value, adapted to

Improved seed size, high fertility, improved germination,

large growth form

Excellent forage attributes, erect, leafy ecotype

Dwarf, blue green, densely tufted, drought tolerant, persistent, competitive

Improved seedling vigor and yields

cold tolerant

NRCS, Pullman, WA Kansas AES Leafy, vigorous, spreads rapidly, competitive

Alaska AES Winter hardy, high yields, early spring growth

Iowa AES Very productive, well-suited to wet sites, useful for hay

U of Idaho, Moscow, ID

Post Falls, ID

Oregon AES Remains green throughout the summer, high forage yields,

Oregon AES High forage and seed production, widely adapted

and smooth brome sources

foreign introductions

Lincoln 1942 Hungary Nebraska AES Aggressive sodformer, good seedling vigor, drought tolerant

Numerous other smooth brome cultivars of both the northern and southern types have been developed for regional uses.

Ioreed 1946 Composite clones

Other less common cultivars include Auburn, Castor, Frontier, Highland, Rise, and Vantage.

Durar 1949 Union, OR, NRCS, Pullman, WA

Nezpurs 1983 Composite clones

MX86 1989 Germany Jacklin Seed Company,

Fawn 1964 Named varieties and

germplasm date Origin seed maintenance

var. 1934 planting semiarid sites

var.

(Con.)

)

ovina

F. ovina

Brome, smooth Manchar 1943 Manchuria, China Common and Cultivar or ReleaseSouthern types Achenbach 1944 Washington County, KS Breeder scientific names

Northern types Polar 1965 Arctic brome hybrids

Canarygrass, reed

Table 1

Fescue, Arizona Redondo 1973 Los Alamos, NM NRCS, Los Lunas, NM

Fescue, hard sheep

Fescue, Idaho Joseph 1983 Composite clones

(

Fescue, sheep Covar 1977 Konya, Turkey NRCS, Pullman, WA var. Bromus inermis Phalaris arundinacea Festuca arizonica Festuca ovina duriuscula Festuca idahoensis ingrata Festuca ovina Fescue, tallFestuca arundinacea Alta 1940 Composite clones

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 299 Chapter 18 Grasses

(con.)

Attributes

Highly productive, excellent quality forage,

heavy seed producer, rhizomatous, adapted to mountain meadows

Improved seedling vigor, high seed production

Intermediate growth habit, fine stemmed

High seed and herbage production, drought tolerant

Excellent seedling vigor, rapid establishment

Outstanding leafiness, forage production, vigor

Strong rhizome production

Outstanding vigor, tolerates cold and dry climates

No rhizomes, matures evenly

Outstanding vigor, leafiness, good seedling vigor

Good seedling vigor, drought tolerant, easily established

Ontario, Canada

NRCS, Los Lunas, NM

NRCS, Bismarck, ND

NRCS, Los Lunas, NM

Nebraska AES Winter hardy, long lived, late maturing

Counties, NE

Counties, TX

Garrison 1963 Europe NRCS, Bismarck, ND

Dan 1987 Wilno, Poland AC, Ottawa, Early maturity, good winter hardiness

Lovington 1963 Lovington, NM NRCS, Los Lunas, NM

Butte 1958 Holt and Platte Nebraska AES Good seedling vigor, winter hardy, long lived

El Reno 1944 El Reno, OK NRCS, Manhatten, KS

Haskell 1983 Haskell, TX NRCS, Knox City, TX

Killdeer 1960 Dunn and Bowman

Niner 1984 Socorro County, NM

Pierre 1960 Pierre, SD NRCS, Bismarck, ND

Premier 1960 Mexico Texas AES Upright, leafy type, seedlings develop rapidly

Trailway 1958 Holt County, NE

Vaughn 1940 Vaughn, NM NRCS, Los Lunas, NM

germplasm date Origin seed maintenance

(Con.)

Foxtail, creeping (reed)

Common and Cultivar or Release Breeder scientific names

Table 1 Foxtail, meadow

Galleta Viva 1979 New Kirk, NM NRCS, Los Lunas, NM

Grama, black Nogal 1971 Socorro County, NM

Grama, blue Hachita 1980 Hachita, NM NRCS, Los Lunas, NM Alopecurus arundinaceus Alopecurus pratensis Hilaria jamesii Bouteloua eriopoda Bouteloua gracilis Grama, sideoates Bouteloua curtipendula

300 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

(con.)

palatable

Attributes

Adapted to high elevations and acidic soils

Upright growth form, leafy, free of galls

High palatability, good regrowth

Late maturing, abundant leaves, highly productive

Drought hardy, easily established, moderately productive

Highly productive, persistent, leafy type

Winter-hardy, robust, cool-season perennial developed

for Alaska

Excellent seedling vigor, low-percentage hard seeds

Soil stabilization, drought tolerant, highly

Good seed retention, low shattering, large stature

Good seedling vigor and forage production

Highly palatable, good germination

Alaska AES Low-maintenance ground cover

NRCS, Meeker, CO

Alaska and Iceland

collections Improvement Association

Norcoast 1981 Alaska Alaska AES High forage production, adapted to semiwet and acid soils

Nortran 1986 Composite collection,

Peru Creek 1995 Summit County, CO

Lodorm 1970 Bismarck, ND ARS, Mandan, ND

Paiute 1983 Ankara, Turkey NRCS, Aberdeen, ID

Potomac 1954 Combined Washington Crop

Numerous other pasture cultivars are available

Sourdough 1976 Alaska NRCS, Palmer, AK

Nezpar 1978 Whitebird, ID NRCS, Aberdeen, ID

Paloma 1974 Pueblo, CO NRCS, Los Lunas, NM

Rimrock 1996 Billings, MT NRCS, Bridger, MT

Saltalk 1981 Erick, OK NRCS, Knox City, TX

Sand Hollow 1996 Gem County, ID Self-pollinating, grows at low temperatures

germplasm date Origin seed maintenance

(Con.)

anadensis

Hairgrass, tufted

Common and Cultivar or Release Breeder scientific names

Muhly, spike El Vado 1973 Park View, NM NRCS, Los Lunas, NM

Needlegrass, green

Orchardgrass Latar 1957 U.S.S.R. NRCS, Pullman, WA Table 1

Reedgrass, bluejoint

c Ricegrass, Indian

Sacaton, alkali Salado 1982 Claunch, NM NRCS, Los Lunas, NM Deschampsia caespitosa Muhlenbergia wrighti Stipa viridula Dactylis glomerata Calamagrostis Oryzopsis hymenoides Sporobolus airoides Squirreltail, bottlebrush Sitanion hystrix

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 301 Chapter 18 Grasses

(con.)

Good seedling vigor,

Attributes

establishes quickly

Shade tolerant, provides late-season forage

Long-lived, highly productive, drought resistant

Excellent seedling vigor and summer greenness recommended for semiarid sites

Good seedling vigor, adapted to semiarid sites

Low growing, slightly rhizomatous, drought tolerant

McGill University, Montreal,Quebec, Canada

NRCS, Pullman, WA

Breeder, seed USDA FS, Ricky Mountain Research Station,

maintenance Provo, UT; NRCS , Aberdeen, ID NRCS, Aberdeen, ID

NRCS, Aberdeen, ID

ARS, Logan, UT Recommended for mesic to semiarid sites

Saskatchewan, Canada

AC, Saskatoon, High seed and forage yields, leafiness, increased vigor Saskatchewan, Canada

progeny Ontario, Canada

of 25 populations

and Nebraska AES

hybrid

Drummond 1940 Northern Europe

Numerous pasture cultivars are available.

Whitmar 1946 Whitman County, WA

Anatone 2003 Asotin County, WA

Douglas 1994 U.S.S.R./Iran/Turkey

Goldar 1989 Asotin County, WA

P-7 2001 Multiple origin polycross

Ephraim 1983 Ankara, Turkey NRCS, Aberdeen, ID

Fairway 1927 U.S.S.R. AC, Saskatoon, Productive, leafy types

NewHy 1989 Quackgrass and ARS, Logan, UT Vigorous, highly productive, high salinity tolerance Parkway 1969 Fairway germplasm

Ruff 1974 Commercial lots Nebraska AES Early, cool season, adapted to low rainfall areas

germplasm date Origin seed maintenance

)

ssp.

)

x bluebunch wheatgrass

inermis

(Con.)

ssp.

Pseudoroegneria Elytrigia repens

Timothy Climax 1947 Combined AC, Ottawa, Highly productive, very leafy,

Common and Cultivar or Release Breeder scientific names

Wheatgrass, beardless (

Table 1

Wheatgrass, bluebunch

Wheatgrass, fairway crested

Wheatgrass, hybrid Phleum pratense Agropyron spicatum inerme spicata Agropyron spicatum, Pseudoroegneria spicata Agropyron cristatum Agropyron repens A. spicatum ( Pseudoroegneria spicata

302 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

(con.)

Attributes

Excellent seedling vigor and early maturity, highly competitive with annuals

Excellent forage traits, good establishment attributes

Drought tolerant, winter hardy, high seed yields

Vigorous, highly productive, late maturing, good regrowth

Vigorous, rhizomatous, drought tolerant

Late maturation, high forage production

Excellent seedling vigor, highly productive, good ground cover

Drought tolerant, dwarf strain, late maturing, sodforming

ARS, Logan, UT Leafy, highly productive, adapted to arid sites

AC, Saskatoon, Improved vigor, fertility, seed development, reduced awns Saskatchewan, Canada

NRCS, Los Lunas, NM

Saskatchewan, Canada

AC, Swift Current, Saskatchewan, Canada

ARS, Mandan, ND

selection from Ree

Ree

and Nebraska 50

SL1 1991 Hybrid ARS, Logan, UT Large culms, productive, improved seedling vigor

CDII 1996 10-clone synthetic

Hycrest 1984 Central Asia ARS, Logan, UT

Kirk 1987 Tetraploid selection

Amur 1952 Manchuria, China

Chief 1961 U.S.S.R. and AC, Saskatoon, Improved forage and seed yields

Clarke 1980 20-clone synthetic

Greenar 1945 U.S.S.R. NRCS, Pullman, WA

Oahe 1961 Selection from South Dakota AES

Reliant 1991 6-clone synthetic

Rush 1995 Germany NRCS, Aberdeen, ID

Slate 1969 Blend of Amur Nebraska AES Erect culms, strongly spreading

Tegmar 1968 Turkey NRCS, Aberdeen, ID

germplasm date Origin seed maintenance

) cristatum

(Con.)

Thinopyrum

Wheatgrass, hybrid Common and Cultivar or Release Breeder scientific names

Wheatgrass, hybrid crested derived from Hycrest

Table 1

Wheatgrass, intermediate Agropyron spicatum A. dasystachyum Agropyron A. desertorum Agropyron intermedium ( intermedium

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 303 Chapter 18 Grasses

(con.)

Attributes

Adapted to warm temperatures and dry sites, excellent seedling vigor, highly productive

Vigorous, drought tolerant, sodformer

Drought resistant, good seedling vigor, persistent

Excellent seedling vigor, highly productive

Highly productive, adapted to saline soils

Vigorous, early growing, long-lived, high yield, alkali tolerant

Tolerant of drought and salinity, vigorous seedlings

Excellent establishment, high seed and forage yields

Excellent establishment and longevity, adapted to high elevations

Dryland ecotype, drought tolerant, tufted bunchgrass

Good seedling vigor, large seeds, high yields, cold hardy

NRCS, Bridger, MT

NRCS, Meeker, CO

Saskatchewan, Canada

seed lots Alberta, Canada winter hardy

Canada Saskatchewan, Saskatchewan, Canada

Saskatchewan, Saskatchewan, Canada Canada

Greenleaf 1966 Commercial AC, Lethbridge, Sod forming, high seedling vigor and forage production,

Luna 1963 U.S.S.R. NRCS, Los Lunas, NM

Topar 1953 U.S.S.R. NRCS, Aberdeen, ID

P27 1953 U.S.S.R. NRCS, Aberdeen, ID

Vavilov 1994 U.S.S.R. NRCS, Aberdeen, ID

Adanac 1990 Climax, AC, Saskatoon,

Primar 1946 Beebe, MT NRCS, Pullman, WA

Pryor 1988 South-central MT

Revenue 1970 Revenue, AC, Saskatoon,

San Luis 1984 Rio Grande County, CO

Secar 1980 Lewiston, ID NRCS, Pullman, WA

Nordan 1953 Central Asia ARS, Mandan, ND

Summit 1953 U.S.S.R. AC, Saskatoon, Excellent forage traits, high seed yields

germplasm date Origin seed maintenance

)

(Con.)

wawawai

Thinopyrum intermedium, A. fragile Elymus trachycaulus Agropyron spicatum,

Wheatgrass, pubescent (

Common and Cultivar or Release Breeder scientific names

Wheatgrass, Siberian

Table 1

(

Wheatgrass, slender (

Wheatgrass, Snake River

(

spp.

Wheatgrass, standard Agropyron intermedium A. trichophorum Agropyron sibiricum Agropyron trachycaulum Elymus wawawaiensis E. lanceolatus crested Agropyron desertorum

304 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

(con.)

Attributes

Tall, robust type, tolerant of wet and saline soils

Drought and salt tolerant, early maturing, fine leaves

Highly productive, large bunch type

Winter hardy, withstands flooding, good seed and forage yield

tolerant of alkali soils

Rhizomatous, highly productive, drought tolerant

Excellent seedling vigor, strongly rhizomatous

Weakly rhizomatous, adapted to dry sites

Drought resistant, strongly rhizomatous, excellent ground cover

Vigorous sodformer and controls weeds

Rapid germination, excellent establishment, good seed and forage yields

Strongly rhizomatous, leafy, highly productive

Spreading type, early spring growth, drought tolerant

Highly productive, excellent pasture type, vigorous

Excellent seedling vigor, close sod, leafy

Nebraska AES Winter hardy bunchgrass with short, spreading rhizomes,

NRCS, Aberdeen, ID

AES, Lincoln, NE

local strain Saskatchewan, Canada

Saskatchewan Alberta, Canada tolerant of flooding and salinity

ritana 1971 Havre, MT NRCS, Bridger, MT

Alkar 1951 U.S.S.R. NRCS, Pullman, WA

Jose 1965 Australia NRCS, Los Lunas, NM

Largo 1961 Turkey NRCS, Los Lunas, NM

Orbit 1966 PI 98526 and AC, Swift Current,

Platte 1972 Nebraska 98526

Bannock 1995 Composite NRCS, Aberdeen, ID

Schwendimar 1994 The Dalles, OR NRCS, Pullman, WA

Sodar 1954 Grant County, MT

Thickspike 1994 The Dalles, OR NRCS, Aberdeen, ID

Arriba 1973 Flagler, CO NRCS, Los Lunas, NM

Barton 1970 Heizer, KS NRCS, Manhattan, KS

Flintlock 1975 Nebraska and Kansas

Rodan 1983 Mandan, ND ARS, Mandan, ND

Rosana 1972 Forsyth, MT NRCS, Bridger, MT

Walsh 1982 Alberta and AC, Lethbridge, Drought tolerant, dense sodformer, high yields,

germplasm date Origin seed maintenance

)

(Con.)

lanceolatus riparium

Thinopyrum ponticum Elymus lanceolatus . Elytrigia smithi

Wheatgrass, tall

Common and Cultivar or( Release Breeder scientific names

Table 1

Wheatgrass, thickspike Agropyron dasystachyum ( var.

Wheatgrass, western Agropyron elongatum A Agropyron smithii ( Pascopyrum smithii

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 305 Chapter 18 Grasses

(con.)

Attributes

Highly productive, good-quality late-season herbage, good

seedling vigor

Highly productive, good production in late summer, good seedling vigor, drought tolerant

Excellent producer, good-quality late-season herbage, free of leaf spot

Excellent seed viability, vigorous spreader or creeper Leafy, fine stemmed, highly productive, rhizomatous

Vigorous, disease-free, high-quality seed

Easily established, short stature, high seed and

forage production, leafy, persistent

Short lived, highly productive first 13 years of stand

establishment, erect, green foliage

Highly productive, good stand establishment

Large, robust, extremely productive, excellent

Vigorous, drought tolerant, early spring forage

Tall, persistent, useful for dune stabilization

Saskatchewan, Canada

Canada ground cover, good seed quality

Pearl 1989 U.S.S.R. AC, Swift Current,

Prairieland 1976 U.S.S.R. AC, Swift Current,

RioShoshone 1980 1991 Riverton, WY Stratford, CA NRCS, Bridger, MT NRCS, Lockford, CA

Mandan 1946 Mandan, ND ARS, Mandan, ND

Arthur 1989 China AC, Swift Current,

James 1989 China AC, Indian Head,

Magnar 1979 Saskatchewan, NRCS, Aberdeen, ID

Trailhead 1991 Roundup, MT NRCS, Bridger, MT

Volga 1949 U.S.S.R. NRCS, Meeker, CO

germplasm date Origin seed maintenance

)

(Con.)

Leymus angustus Leymus triticoides Leymus cinereus Leymus racemosus

Wildrye, Altai Eejay 1989 U.S.S.R. AC, Indian Head,

Common and Cultivar or Release Breeder scientific names

(

Table 1

Wildrye, beardless (

Wildrye, blue Arlington 1995 Arlington, WA NRCS, Corvallis, OR

Wildrye, Canada

Wildrye, Dahurian

( Wildrye, Great Basin

( Elymus angustus Elymus triticoides Elymus glaucus Elymus canadensis Elymus dahuricus Elymus cinereus Wildrye, mammoth Elymus giganteus

306 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Attributes

on; AC = Agriculture Canada; ARS = USDA Agricultural

Improved seed retention, seedling vigor, and forage

Productive, good late-summer herbage, resistant to leaf spot disease

Improved seedling vigor, highly productive, good seed quality and yield

Large seeds, improved seedling vigor

Provides early greenup and fall grazing, high seed

Alberta, Canada and seed yield

Mandan, ND Excellent fall and winter grazing, seedling establishment,

AC, Swift Current,

Manyberries, Alberta, Canada seeding

U.S.S.R. and and forage production Mandan, ND

synthetic Saskatchewan, Canada

tetraploids Saskatchewan, Canada

synthetic production

Select Cabree 1976 Six clones from AC, Lethbridge,

Mankota 1991 Six clone synthetic,

Mayak 1971 Twenty-clone AC, Swift Current,

Swift 1978 Twenty-six clone

Tetracan 1988 Colchicine induced

Vinall 1960 Five clone ARS, Mandan, ND

germplasm date Origin seed maintenance

(Con.)

)

NRCS = USDA Natural Resource Conservation Service, formerly USDA Soil Conservation Service; AES = Agricultural Experiment Stati

a Psathyrostachys juncea

Table 1

Common andWildrye, Russian Cultivar or Release Bozoisky- 1984 U.S.S.R. ARS, Logan, UT Breeder Improved seedling vigor, highly productive scientific names

( Elymus junceus Research Service; UCEPC = Upper Colorado Environmental Plant Center.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 307 Chapter 18 Grasses

Table 2—Characteristics of selected grasses.

Species

Seed production Ease of seed harvesting Ease of seed cleaning Ease of seeding Germination Seedling vigor Growth rate Final establishment Method of spread Spring growth Summer growth Fall growth Compatibility with other species Longevity Forage production Spring palatability Summer palatability Grazing tolerance Soil stabilization Shade tolerance Flood tolerance Ecotypic variability

Vegetative types

------ratings b------Barley, bulbous 4 4 3 4 5 5 4 2 S 5 2 2 3 4 3 5 5 5 3 2 2 4 WM,JP,BS Barley, meadow 3 4 3 4 4 5 4 3 V 5 4 3 4 3 3 5 4 3 3 4 4 2 SA,A,WM Bluegrass, big 3 5 5 4 5 5 4 4 S 4 3 4 4 4 3 4 4 4 4 4 3 3 A,MB,JP,PP,MS,R Bluegrass, Canada 3 5 5 4 4 5 4 4 V 5 3 4 4 4 3 4 4 4 4 4 2 3 SA,A,MB,PP Bluegrass, Kentucky 3 5 5 4 4 5 4 5 V 5 4 4 2 5 2 4 4 5 5 4 4 5 SA,A,WM,MB,PP, JP,MS,R Bluegrass, mutton 2 3 5 4 4 5 3 5 S 5 5 4 4 4 4 4 4 3 3 3 3 3 MB,PP,JP,MS Bluegrass, Sandberg 3 5 5 4 4 4 4 5 S 5 2 3 4 5 2 5 2 4 4 2 2 5 MB,PP,JP,MS, BS,WS,SS,C,AW Brome, mountain 4 5 5 4 5 5 5 4 S 3 4 4 5 3 4 4 4 4 4 4 4 3 SA,A,MB,PP,MS Brome, nodding 4 5 5 4 5 5 5 4 S 4 4 4 5 3 4 4 4 4 4 5 4 3 SA,A,MB,PP,MS Brome, smooth 5 5 5 4 5 5 4 5 V 4 4 4 1 5 5 4 4 5 5 5 4 4 SA,A,WM,MB, PP,MS,R northern Brome, smooth 5 5 5 4 5 5 5 5 V 4 4 3 1 5 4 4 3 5 5 3 4 4 SA,A,WM,MB, southern PP,JP,MS,BS,R Brome, subalpine 3 5 5 4 5 4 3 4 S 4 4 4 4 4 4 4 4 3 4 5 4 3 SA,A,WM,MS Canarygrass, reed 5 3 4 4 3 2 5 5 V 3 5 3 1 5 5 3 3 5 5 2 5 2 SA,A,WM,IS,R Dropseed, sand 3 3 4 4 3 2 4 4 S 5 3 2 4 4 3 3 2 4 4 2 3 3 MB,PP,JP,BS, WS,SS,BB,C,AW Fescue, hard sheep 4 5 5 5 5 4 4 5 S 4 3 2 1 5 2 2 2 5 4 2 2 4 A,MB,PP,JP,MS,BS Fescue, Idaho 3 5 5 5 5 4 3 3 S 5 3 2 4 4 3 5 4 4 4 2 2 3 MB,PP,MS Fescue, meadow 3 5 5 5 5 4 4 5 S 4 4 3 2 4 4 5 4 5 5 3 4 2 SA,A,WM,JP,BG,IS Fescue, sheep 3 5 5 5 5 4 4 4 S 5 3 2 3 4 3 3 2 4 4 2 2 5 SA,A,MB,PP,MS Fescue, spike 2 4 4 4 3 3 4 3 V 4 3 2 4 4 4 3 3 4 3 3 3 2 MB,PP,JP,MS Fescue, reed 5 3 3 3 5 4 4 5 S 4 4 3 2 2 4 5 4 4 5 3 5 2 SA,A,WM,JP,BG,IS Fescue, Thurber 2 3 4 4 4 4 3 3 S 5 2 2 4 5 3 4 3 4 3 2 2 4 SA,A,MB,MS Foxtail, meadow 5 2 2 2 4 2 4 5 V 4 4 4 2 5 5 5 5 5 5 3 5 2 SA,A,WM,MB,IS Foxtail, creeping 5 2 2 2 4 2 4 5 V 4 4 4 2 5 5 5 5 5 5 3 5 2 SA,A,WM,MB,IS Galleta 3 3 3 3 3 4 3 3 V 3 4 2 4 4 3 4 4 3 4 2 1 5 JP,BS,WS,SS,BB Grama, sideoats 3 4 4 5 4 4 3 3 S 3 4 2 4 4 3 4 2 3 4 2 1 3 JP,BS,WS,SS,BB Grama, blue 2 4 4 5 4 4 3 4 V 3 4 2 3 5 2 4 2 5 5 3 1 3 JP,BS,WS,SS,BB Grama, black 2 4 4 5 4 4 3 4 V 3 4 2 3 5 2 4 2 5 5 3 1 3 JP,BS,WS,SS,BB Hair grass, tufted 3 4 4 4 4 4 4 4 S 3 4 2 4 4 2 2 2 4 3 2 2 3 SA,A,WM,MB,PP,IS Junegrass, prairie 2 4 5 4 3 4 4 3 S 5 3 3 5 4 2 5 3 4 3 3 2 3 MB,PP,JP,MS, JP,WS,BB Needle-and-thread 3 2 2 3 3 4 3 5 S 5 4 3 3 5 3 4 2 4 4 2 2 3 MB,PP,JP,BS, WS,C,AW Needlegrass, green 4 3 2 3 4 4 4 4 S 5 4 3 4 4 4 4 3 4 4 3 3 3 A,MB,PP,MS Needlegrass, Letterman 3 3 2 3 3 3 3 4 S 3 5 2 4 5 2 2 2 4 4 3 3 2 SA,A,MB,MS Needlegrass, subalpine 2 2 2 3 3 3 3 3 S 3 5 3 4 5 2 4 3 4 4 4 3 2 SA,A,MS Needlegrass, Thurber 3 2 2 3 2 3 3 5 S 5 4 3 3 5 4 4 4 4 4 3 2 3 MS,BS,WS,C,AW Oatgrass, tall 4 3 3 4 5 4 4 4 S 5 4 3 4 3 4 5 4 3 4 5 3 2 SA,A,MB,PP,MS Oniongrass 2 5 5 5 5 4 5 3 V 4 3 3 4 2 3 4 4 3 3 5 3 2 A,MB,PP,MS Orchardgrass 5 5 5 5 5 5 5 4 S 5 4 4 3 4 5 5 4 4 5 5 4 5 SA,A,WM,MB, PP,MS Redtop 3 3 4 4 3 3 3 5 V 3 4 2 3 4 2 3 2 4 3 3 4 3 A,WM,BS,BG,IS Ricegrass, Indian 4 3 3 4 2 4 3 4 S 5 2 2 4 5 3 4 3 4 4 2 5 4 MB,PP,PJ,MS,BS, WS,SS,BG,BB,C,AW (con.)

308 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Table 2 (Con.)

Species

Vegetative types

------ratings b------Rye, mountain 5 4 4 5 5 5 5 4 S 5 2 5 3 2 5 5 4 4 4 3 2 3 MB,PP,JP,MS,BS,WS,SS, BB,C,AW Sacaton, alkali 3 3 4 4 1 2 2 4 V 3 3 3 1 5 2 2 2 5 5 2 4 3 WM,BS,WS,SS,BG, BB,IS,R Squirreltail, bottlebrush 4 2 2 3 4 4 4 4 S 5 2 2 4 4 3 4 2 4 4 2 5 5 MB,PP,JP,BS,WS,SS,BG, BB,C,AW Three-awn, purple 2 2 2 3 3 4 3 4 S 5 2 2 3 5 3 1 1 5 4 2 3 4 MB,PP,JP,MB,BS, WS,SS,C,AW Timothy, mountain 4 4 4 4 5 4 5 4 S 3 4 3 4 3 3 5 5 5 4 4 5 3 SA,A,WM Timothy 5 4 4 4 5 5 5 5 S 4 4 4 4 3 5 5 5 4 3 4 4 3 SA,A,WM,MB,PP,JP,MS Trisetum, spike 2 2 4 3 4 3 4 3 S 5 4 3 4 3 2 3 3 3 3 3 2 3 MB,JP,MS,BS,WS Wheatgrass, bluebunch 5 5 5 4 5 4 4 4 S 5 3 4 4 5 5 5 4 4 4 3 5 5 MB,PP,JP,MS,BS, WS,C,AW Wheatgrass, fairway 5 5 5 5 5 5 5 5 S 5 3 3 1 5 4 5 3 5 5 3 4 3 MB,PP,JP,MS,BS,WS,SS, crested BG,BB,C,AW Wheatgrass, 5 5 5 5 5 5 5 5 V 5 4 3 1 5 5 5 4 5 5 4 4 5 A,MB,PP,JP,MS,BS,WS, intermediate SS,R,C,AW Wheatgrass, pubescent 5 5 5 5 5 5 5 5 V 4 3 5 1 5 5 5 3 5 5 2 2 5 MB,PP,JP,MS,BS,WS, SS,BB,R,C,AW Wheatgrass, Siberian 3 5 5 5 5 4 4 4 S 5 2 2 2 5 3 5 3 5 4 2 2 4 BS,WS,SS,BG,BB, crested IS,C,AW Wheatgrass, slender 4 5 5 5 4 5 4 4 S 4 4 3 4 3 4 5 4 4 4 4 3 5 SA,A,MB,PP,JP,MS Wheatgrass, Snake River 5 5 5 5 5 5 4 4 S 5 3 4 4 5 5 5 4 4 4 2 2 5 MB,PP,JP,MS,BS, WS,C,AW Wheatgrass, 5 5 5 5 5 5 5 5 S 5 2 3 1 5 4 5 3 5 5 3 2 4 MB,PP,JP,MS,BS,WS, standard crested SS,BG,BB,C,AW Wheatgrass, tall 5 5 5 5 5 5 5 4 S 4 3 3 2 4 5 3 2 4 4 3 5 4 MB,PP,JP,MS,BS,WS, BG,IS,R,C,AW Wheatgrass, thickspike 4 5 5 5 4 4 4 3 V 5 3 3 2 5 3 4 2 5 5 4 3 5 MB,PP,JP,MS,BS Wheatgrass, western 3 5 5 5 3 3 3 5 V 5 3 3 3 5 3 4 3 5 5 4 3 5 MP,PP,JP,MS,BS,WS, SS,BG,R,C,AW Wildrye, Altai 5 5 5 4 2 2 3 5 V 4 4 3 3 4 5 4 3 5 5 4 4 3 MB,PP,JP,BG,IS,R Wildrye, blue 4 5 5 4 5 5 4 3 S 4 4 3 3 4 5 4 4 2 4 3 3 2 A,MB Wildrye, Canada 4 3 2 4 4 3 4 4 S 4 4 3 4 4 4 4 4 3 4 4 3 2 A,MB,WM,IS,R Wildrye, alkali 3 4 5 4 4 4 4 4 V 4 4 2 3 4 4 4 3 4 5 2 3 2 WM,JP,BS,WS,SS,BG,IS Wildrye, Great Basin 5 4 5 4 4 3 3 3 S 5 4 2 4 5 5 4 3 2 4 2 4 5 WM,MB,MS,BG,IS, R,C,AW Wildrye, mammoth 1 3 5 4 3 3 3 2 V 4 4 2 4 5 5 2 2 4 4 2 4 4 MB,JP,MS,BS Wildrye, Russian 4 4 5 5 4 3 3 5 S 5 4 4 4 5 4 5 5 5 4 2 2 4 MS,JP,MS,BS,WS,SS, BG,BB,IS,C,AW Wildrye, Salina 2 4 5 4 1 2 2 4 V 4 3 2 2 5 2 1 1 5 5 2 3 2 MB,PP,JP,MS,BS,WS, SS,BG,IS,C,AW

aVegetative types to which the species is adapted: A = Aspen-conifer; AW = Annual weeds; BB = Blackbrush; BG = Black greasewood; BS = Basin big sagebrush; C = Cheatgrass; IS = Inland saltgrass; JP = Juniper-pinyon; MB = Mountain brush; MS = Mountain big sagebrush; PP = Ponderosa pine; R = Riparian; SA = Subalpine; SS = Shadscale saltbush; WM = Wet and semiwet meadows; WS = Wyoming big sagebrush. b Key to ratings: 1 = Poor Ð difficult; 2 = Fair; 3 = Medium; 4= Good; 5 = Excellent Ð easy; A = Annual - reproduction from seed; S = Reproduction from seed; V = Reproduction vegetative and from seed .

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 309 Chapter 18 Grasses

(con.)

USDA NRCS (2002)

Wheatgrass, crested Wheatgrass, streambank

Wheatgrass, rush Wheatgrass, beardless Wheatgrass, intermediate Wheatgrass, Russian

Wheatgrass, steambank Wheatgrass, spreading Wheatgrass, Siberian

Wheatgrass, western Wheatgrass, bluebunch Wheatgrass, slender Wheatgrass, slender Wheatgrass, intermediate Redtop

Bentgrass, creeping

Brome, meadow Brome, California

Brome, mountain Brome, Pumpelly

Cheatgrass Bluejoint Reedgrass, chee

Hairgrass, tufted

Wildrye, Canada Wildrye, basin Wildrye, mammoth

Wildrye, Russian Wildrye, mammoth

Common name

Plummer and others (1977)

Wheatgrass, fairway crested Wheatgrass, thickspike Wheatgrass, standard crested (desert) Wheatgrass, desert Wheatgrass, tall Wheatgrass, beardless bluebunch Wheatgrass, intermediate Wheatgrass, rushleaf Quackgrass Quackgrass

Wheatgrass, steambank Wheatgrass, Scribner Wheatgrass, Siberian

Wheatgrass, western Wheatgrass, bluebunch Wheatgrass, bearded Wheatgrass, slender Wheatgrass, pubescent Bentgrass, redtop

Bentgrass, carpet Foxtail, reed Foxtail, creeping meadow Foxtail, meadow Foxtail, meadow Threeawn, red Threeawn, fendler Oatgrass, tall Oatgrass, tall Grama, sideoats Grama, sideoats Grama, blue Grama, blue Brome, nodding Brome, nodding

Brome, Regar (meadow) Brome, mountain

Brome, meadow Brome, erect Brome, smooth Brome, smooth Brome, big mountain Brome, Pumpelly Brome, foxtail Brome, red

Brome, cheatgrass Reedgrass, bluejoint Reedgrass, chee Orchardgrass Orchardgrass Hair-grass, tufted Saltgrass, inland Saltgrass, inland Wildrye, Altai Wildrye, Altai Wildrye, purple Wildrye, purple

Wildrye, Canada Wildrye, Great Basin Wildrye, yellow Wildrye, yellow Wildrye, mammoth

Wildrye, blue Wildrye, blue Wildrye, Russian Wildrye, sabulosa

Wildrye, salina Wildrye, saline

)

spicata inermis

subsecundus trachycaulus

ssp.

ssp. lanceolatus sabulosa

longiseta

ssp. ssp. pumpellianus Leymus triticoides salinus

ssp.

var.

ssp.

Elymus elymoides

Agropyron cristatum Elymus lanceolatus Agropyron desertorum Thinopyrum ponticum Thinopyrum intermedium Thinopyrum junceiforme Elymus lanceolatus Elymus scribneri Agropyron fragile Pseudoroegneria spicata Elymus trachycaulus Elymus trachycaulus Thinopyrum intermedium Agrostis gigantea Agrostis stolonifera Alopecurus arundinaceus Alopecurus pratensis Aristida purpurea Arrhenatherum elatius Bouteloua curtipendula Bouteloua gracilis Bromus anomalus Bromus biebersteinii Bromus carinatus Bromus erectus Bromus marginatus Bromus tectorum Calamagrostis canadensis Calamagrostis epigeios Dactylis glomerata Deschampsia caespitosa Leymus angustus x Elyleymus aristatus ( Elymus canadensis Leymus flavescens Leymus racemosus Leymus racemosus

ND Agropyron cristatum Elymus lanceolatus Agropyron desertorum Elymus elongatus Elymus spicatusElymus hispidus Elymus lanceolatus Pseudoroegreria spicata Elymus scribneri Agropyron cristatum Elymus smithiiElymus spicatus Elymus trachycaulus Elymus trachycaulus Elymus hispidus Agrostis stolonifera Pascopyrum smithii Agrostis stolonifera Alopecurus pratensis Aristida purpurea Arrhenatherum elatius Bouteloua curtipendula Bouteloua gracilis Bromus anomalus Bromus biebersteinii Bromus carinatus Bromus inermisBromus carinatus Bromus inermisBromus rubensBromus tectorum Calamagrostis canadensis Bromus inermis Dactylis glomerata Bromus inermis Deschampsia cespitosa Bromus rubens Distichlis spicataElymus canadensis Distichlis spicata Elymus cinereusND Elymus glaucusElymus junceus Leymus cinereus Elymus glaucus Psathyrostachys juncea

Scientific name

Dewey (1984) Arnow (1987) USDA NRCS (2002)

ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND

ND ND Agropyron cristatum Elymus lanceolatus Agropyron desertorum Thinopyrum elongatum Pseudoroegneria spicata Thinopyrum intermedium Thinopyrum junceiforme repensElytrigia Elymus lanceolatus Elymus scribneri Agropyron fragile Pascopyrum smithii Pseudoroegneria spicata Elymus repensElymus trachycaulus Elymus trachycaulus Thinopyrum intermedium Elymus repens Leymus angustus Elymus canadensis Leymus cinereus Leymus flavescens Leymus racemosus ND ND

Barkworth and Dewey (1985),

—Scientific names, synonyms, and common names of grass species. ND = Species not described in reference.

Hitchcock (1951)

Table 3 Agropyron cristatum Agropyron dasystachyum Agropyron desertorum Agropyron elongatum Agropyron inerme Agropyron intermedium Agropyron junceum Agropyron repens Agropyron riparium Agropyron scribneri Agropyron sibiricum Agropyron smithii Agropyron spicatum Agropyron subsecundum Agropyron trachycaulum Agropyron trichophorum Agrostis alba Agrostis stolonifera Alopecurus arundinaceus Alopecurus pratensis Aristida longiseta Arrhenatherum elatius Bouteloua curtipendula Bouteloua gracilis Bromus anomalus Bromus biebersteinii Bromus carinatus Bromus erectus Bromus inermis Bromus marginatus Bromus pumpellianus Bromus rubens Bromus tectorum Calamagrostis canadensis Calamagrostis epigeios Dactylis glomerata Deschampsia caespitosa Distichlis spicata Elymus angustus Elymus aristatus Elymus canadensis Elymus cinereus Elymus flavescens Elymus giganteus Elymus glaucusElymus junceusElymus sabulosus Elymus salinus Leymus glaucus Psathyrostachys juncea Leymus salinus Elymus salinus Leymus salinus

310 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

USDA NRCS (2002)

Wildrye, alkali Wildrye, beardless

Fescue, tall Ryegrass, meadow

Fescue, hard

Fescue, Thurber

Barley, common

Junegrass, prarie Ryegrass, Italian Ryegrass, perennial

Ricegrass, Indian Canarygrass, reed

Bluegrass, bulbous

Bluegrass, Sandberg Bluegrass, Canada Bluegrass, Cusick Muttongrass

Muttongrass Bluegrass, Sandberg

Bluegrass, Kentucky Bluebrass, nodding

Bluegrass, Sandberg

Squirreltail

Needlegrass, Columbia Needle-and-thread Needlegrass, Letterman Needlegrass, Thurber Neddlegrass, green

Common name

Plummer and others (1977)

Wildrye, low creeping Wildrye, creeping Fescue, Arizona Fescue, Arizona

Fescue, reed (Alta, tall) Fescue, meadow Fescue, Idaho Fescue, Idaho

Fescue, hard sheep

Fescue, sheep Fescue, sheep Fescue, Thurber Fescue, spike Fescue, spike Galleta Galleta, James Barley, meadow Barley, meadow Barley, bulbous Barley, bulbous Barley, foxtail Barley, foxtail Barley, beardless

Junegrass, prarie Ryegrass, Italian Ryegrass, perennial Oniongrass Oniongrass Ricegrass, Indian Canarygrass, reed Timothy, alpine Timothy, alpine Timothy Timothy Bluegrass, big Bluegrass, Sandberg Bluegrass, bulbous

Bluegrass, Canby Bluegrass, Canada Bluegrass, Cusick Bluegrass, mutton

Bluegrass, longtongue Bluegrass, Nevada

Bluegrass, Kentucky Bluegrass, nodding Bluegrass, pine Bluegrass, Sandberg Bluegrass, Sandberg

Squirreltail, bottlebrush Sacaton, alkali Sacaton, alkali Dropseed, sand Dropseed, sand Needlegrass, subalpine Needle-and-thread Needlegrass, Letterman Needlegrass, Thurber Needlegrass, green Trisetum, spike Tristetum, spike

multiflorum longiligula

ssp.

Leymus triticoides Festuca arizonica Lolium arundinaceum Lolium pratense Festuca idahoensis Festuca trachyphylla Festuca thurberi Leucopoa kingii Hordeum brachyantherum Hordeum bulbosum Hordeum jubatum Hordeum vulgare Koeleria macrantha Achnatherum hymenoides Phalaris arundinacea Phleum pratense NDElymus elymoides Sporobolus airoides Sporobolus cryptandrus Achnatherum lettermanii Achnatherum thurberianum Trisetum spicatum Rye, mountain ND

ovina Festuca ovina

var.

ingrata duriuscula

Elymus simplexElymus triticoides Festuca ovina arizonica Festuca arundinacea Festuca pratensis Leymus simplex Festuca ovina Festuca ovina Festuca ovina Festuca thurberi Leucopoa kingii Hilaria jamesiiHordeum brachyantherum Horedum jubatum Horedum vulgare Koeleria macrantha Pleuraphis jamesii Lolium perenneLolium perenneMelica bulbosaStipa hymenoides Phalaris arundinacea Phleum alpinum Lolium perenne Phleum pratense Lolium perenne Poa secunda Melica bulbosa Poa bulbosaPoa secunda Phleum alpinum Poa compressaPoa fendlerianaPoa fendlerianaPoa fendleriana Poa secunda Poa secunda Poa bulbosa Poa pratensis Poa secunda Poa compressa Poa reflexa Poa cusickii Poa secunda Poa fendleriana Poa secunda Poa fendleriana Secale montanum Elymus elymoides Poa secunda Poa pratensis Sporobolus airoides Sporobolus cryptandrus Poa reflexa Stipa nelsonii Poa secunda Stipa comata Poa secunda Stipa lettermanii Stipa thurberiana Stipa viridulaTrisetum spicatum Achnatherum nelsonii Hesperostipa comata Nassela viridula

Scientific name

Dewey (1984) Arnow (1987) USDA NRCS (2002)

ND ND ND

ND ND ND ND

ND ND

ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND

ND ND ND ND ND ND ND Leymus triticoides Hordeum brachyantherum Hordeum jubatum ND

Barkworth and Dewey (1985),

ovina

var. ND var.

(Con.)

Hitchcock (1951)

Table 3 Elymus simplexElymus triticoides Festuca arizonica Festuca arundinacea Festuca elatior Leymus simplex Festuca idahoensis Festuca ovina duriuscula Festuca ovina Festuca thurberi Hesperochloa kingii Hilaria jamesii Hordeum brachyantherum Hordeum bulbosum Hordeum jubatum Hordeum vulgare Koeleria cristata Lolium multiflorum Lolium perenne Melica bulbosa Oryzopsis hymenoides Phalaris arundinacea Phleum alpinum Phleum pratense Poa ampla Poa bulbosa Poa canbyi Poa compressa Poa cusickii Poa fendleriana Poa longiligula Poa nevadensis Poa pratensis Poa reflexa Poa scabrella Poa secunda Secale montanum Sitanion hystrixSporobolus airoides Sporobolus cryptandrus Stipa columbiana Stipa comata Stipa lettermanii Elymus elymoides Stipa thurberiana Stipa viridula Trisetum spicatum

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 311 Chapter 18 Grasses and about 0.04 inch (l mm) long. The spike is 0.8 to 4 Ecological Relationships and Distribution inches (2 to 10 cm) long and 0.2 to 0.9 inch (5 to 23 mm) wide. Spikelets are several times longer than the Problems relative to the proper identification and internodes. Each produces 3 to 10 florets, but the use of the crested wheatgrasses arose, in part, from upper florets are sterile. Glumes and lemmas are the failure of earlier workers to identify them correctly tapered to a short awn about 0.12 inch (3 mm) long. and to recognize their differing ecological require- Arnow (1987), Brown (1979), Cronquist and others ments. In addition, although the name “crested wheat- (1977), Hitchcock and others (1969), Gould (l951), grass” refers to the comblike spike of fairway crested Looman (1982), and Powell (1994) provide detailed wheatgrass, this common name has been applied to all descriptions of the species. three taxa. Dillman (1946) suggested that some of the Although fairway crested wheatgrass resembles stan- confusion resulted from a 1910 report he prepared in dard crested wheatgrass and Siberian wheatgrass, which both fairway crested wheatgrass and standard some differences can usually be observed (fig. 1). The crested wheatgrass were referred to as A. cristatum major taxonomic feature used to differentiate the (Dillman 1910). This was compounded in 1935 when three grasses is the spike shape. Fairway crested Hitchcock (1935) used a drawing of A. desertorum to wheatgrass spikes are shorter, wider at the base, and illustrate A. cristatum in the “Manual of the Grasses more distinctly tapered at the tip than those of stan- of the United States.” dard crested wheatgrass. Spikes of Siberian wheat- To address these problems, Weintraub (1953) ap- grass are relatively long and narrow compared to plied the following common names: crested wheat- those of the other two wheatgrasses. Fairway crested grass = A. cristatum; desert wheatgrass = A. deser- wheatgrass spikelets and awns are divergent and torum; and Siberian wheatgrass = A. sibiricum. Beetle arranged in a “comblike” manner, while those of stan- (1961) recommended acceptance of this usage, but the dard crested wheatgrass and Siberian wheatgrass confusion continued. Dewey (1986) later suggested that grow at a strong upward angle. Compared to fairway the most appropriate names for the three groups in crested wheatgrass, the stems of standard crested North America are fairway crested wheatgrass (A. cris- wheatgrass are more erect and more or less uniform in tatum), standard or desert crested wheatgrass (A. deser- height, its leaves are mostly basal, and the upper leaf torum), and Siberian crested wheatgrass (A. sibiricum) blades are generally without hairs. (fig. 1). Differences between Eurasian and North American taxonomic treatments further compound this situation and have yet to be resolved. The crested wheatgrasses constitute a polyploid complex (Dewey 1986; Taylor and McCoy 1973). Dip- loid (2n = 14) fairway crested wheatgrass is native to the Steppe region of Eastern Russia and Southwest- ern Siberia (Asay 1995a,b; Dewey 1966, 1969, 1973; Dewey and Asay 1982; Konstantinov 1920). Dewey and Asay (1982) identified different morphological growth forms of diploid introductions, but found that they hybridized readily. Tetraploids (2n = 28) include standard and Siberian wheatgrass and are widely distributed from central Europe to the Middle East, Siberia, Mongolia, and China (Asay and Dewey 1979; Dewey 1986; Knowles 1955; Tsvelev 1976). The hexaploids (2n = 42) are less common and more scattered, occurring in Turkey, Iran, and Kazakhstan (Asay and others 1990; Dewey and Asay 1975). All possible crosses have been made among the three ploidy levels (Asay and Dewey 1979; Asay and Knowles 1985a,b; Dewey 1971, 1974), but breeding has generally been confined to selections within ploidy levels, particularly the diploid and tetraploid levels (Asay 1986). Crested wheatgrass was first introduced in the United States by the U.S. Department of Agriculture Figure 1—Spikelets of fairway crested wheatgrass in 1889 (Dillman 1946; Reitz and others 1936). Mate- (center) are divergent and comblike, while those of rials were obtained from the Valuiki Experiment Sta- standard crested wheatgrass (left) and Siberian crested tion, Samara Government, on the Volga River, some wheatgrass (right) grow upward (RMRS photo). 150 miles (240 km) north of St. Petersburg, Russia.

312 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Additional seed was obtained from the Valuiki Station stands produce 250 to 300 pounds per acre (275 to 325 in 1906. This shipment included six seed lots of 4 to 5 kg per ha) (Westover and others 1932). Less than 100 pounds (1.8 to 2.3 kg) each. One lot, S.P.I. 19536, was pounds per acre (110 kg per ha) are normally har- labeled A. cristatum and five lots, S.P.I. 19537 to19541, vested from rangeland stands in the Intermountain A. desertorum. Plantings and research with crested area. Row spacings of 36 to 42 inches (91 to 107 cm) are wheatgrass began in 1908 at the Belle Fourche Sta- recommended for cultivated fields. This equates to 5 to tion, Newell, SD, and in 1915 at the Northern Great 8 pounds of seed per acre (5.5 to 9.5 kg per ha). Plains Field Station in Mandan, ND (Dillman 1946). Planting at 3 to 4 pounds per acre (3.4 to 4.5 kg per ha) Studies initiated in Canada greatly advanced the provides satisfactory, but slower developing, stands development and use of crested wheatgrass (Kirk (Westover and Rogler 1947). Direct combining pro- 1928). Kirk and others (1934) reported that seed vides seed lots with purities of 50 to 80 percent; this obtained from the U.S. Department of Agriculture was may be increased to 95 percent purity by screening and planted in University of Saskatchewan experimental fanning. Germination is often near 90 percent. Seed plots. Additional introductions were obtained from can usually be stored in a warehouse for several years Russia, Siberia, and Mandan, ND. In 1927, seed ob- without loss of viability. tained from Montana was planted at Manyberries, Fairway crested wheatgrass is one of the most reli- Alberta. The University of Saskatchewan developed able plants to establish from drill seedings on adapted two strains. One, identified as ‘Fairway’ (S-10), ulti- range and wildland sites. It generally develops fairly mately gained widespread use (Kirk and others 1934). uniform stands, even on highly variable sites. Maynard Fairway was initially distributed by the Field Hus- and Gates (1963) reported that establishment success bandry Department, University of Saskatchewan, in of crested wheatgrass could be attributed to the toler- 1927, and licensed in 1932 as a variety (Kirk 1932; ance of the seeds to extreme fluctuations in soil mois- Knowles 1956). Dillman (1946) concluded that the ture. Wetting and drying cycles can increase germina- Fairway variety originated from the same accession tion. Plummer (1943) reported that the success of S.P.I. 19536, A. cristatum. crested wheatgrass stems from its ability to produce a Scientists working in Montana recognized that the greater total root length in the seedling stage than erect bunch-type growth forms of A. desertorum dif- other grass species. fered from fairway-type crested wheatgrass. The Mon- Drill seeding rangeland sites with 2 to 16 pounds of tana Station was the first to apply the name “standard fairway crested wheatgrass seed per acre (2.2 to 17.6 crested wheatgrass” for certified seed increased from kg per ha) was initially advised for sites receiving S.P.I. 19537, an A. desertorum accession (Dillman more than 12 inches (310 mm) of precipitation, and 5 1946). The selection was used as a standard of com- to 6 pounds per acre (5.5 to 6.6 kg per ha) for sites parison for assessment of other selections (Dewey receiving less than 12 inches (310 mm) of moisture 1986). The name “standard crested wheatgrass” slowly (Thomas and others 1945). More recently, seeding evolved to distinguish any nonfairway-type crested recommendations have been reduced to 2 to 4 pounds wheatgrass. Standard is not a specific cultivar, but is per acre (2.2 to 4.4 kg per ha) for either drilling or now used to identify all standard-type A. desertorum. broadcast seeding. When fairway crested wheatgrass Seed of standard wheatgrass was first produced com- is seeded alone in areas invaded by weeds, the rate mercially by a grower in Montana shortly after 1920, should be increased 2 to 3 times. Seeding in dense and was first listed in 1927 by a dealer in North annual weeds, however, is not advisable. Springfield Dakota (Westover and others 1932). (1965) reported that crested wheatgrass stands will reach an equilibrium within 5 to 8 years, regardless of Plant Culture the amount of seed sown or the drill row spacing used. He found similar herbage yields on sites seeded at 2, 4, Good seed crops of fairway crested wheatgrass are and 6 pounds seed per acre (2.2, 4.4, and 6.6 kg per ha) regularly produced on irrigated farmlands in Western at 5-, 12-, and 18-inch (13-, 30-, 46-cm) row spacings in Canada, the Pacific Northwest, the Intermountain the fifth, sixth, and eighth years after seeding. area, and the Great Plains (Hafenrichter and others Fairway crested wheatgrass seed should be drilled 1949; Knowles 1956; Reitz and others 1936; Westover or otherwise planted at depths of 0.5 to 1.0 inch (1.3 to and Rogler 1947; Westover and others 1932). In these 2.5 cm). Planting at greater depths in heavy-textured areas, dryland farms, pastures, and rangeland soils is not advised. Compared with many other range- plantings are also managed for seed production. Seed land grasses, fairway crested wheatgrass establishes is harvested from these plantings only during years better with limited seedbed preparation. McGinnies with adequate moisture, but large quantities of seed (1962), however, found that packing to create a firm may be harvested in good precipitation years. Yields seedbed improved planting success. Cultipacking be- from irrigated fields average 300 to 600 pounds per fore seeding provides a firm seedbed, permitting fur- acre (325 to 675 kg per ha), while yields from dryland row openers to function more effectively. Broadcast

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 313 Chapter 18 Grasses seeding without some means of seed coverage is not more than standard crested wheatgrass (Westover recommended. Direct drilling or broadcast seeding and Rogler 1947). In the United States, the Civilian and covering the seed following burning or removal of Conservation Corps used both fairway and standard competition is generally successful. Stevens (1987b) crested wheatgrass extensively for conservation reported that fairway crested wheatgrass plants es- plantings in the 1930s. Use increased in 1945 follow- tablish extremely well following aerial seeding and ing a special Congressional Appropriation for research chaining on pinyon-juniper and big sagebrush distur- to improve rangelands (Reynolds and Springfield 1953). bances. Fairway crested wheatgrass is well adapted to the Fall seedings are recommended, but spring plantings Northern and Central Great Plains and the drier on prepared seedbeds may be successful if adequate communities of the Intermountain region. Fairway spring precipitation is received. Frischknecht (1951) and standard crested wheatgrass are seeded to stabi- found that the total emergence from crested wheat- lize marginal farmlands and deteriorated rangelands grass and other grass seedings was greater from early in the Western United States. Fairway crested wheat- fall than from late fall or spring plantings, and that grass has been used in mixtures with other grasses cold temperatures accounted for most grass seedling and alfalfa for hay production. It has also been planted mortality. Buman and Abernethy (1988) found that alone in lawns, yards, and on roadways where supple- mountain rye, crested wheatgrass, and cheatgrass mental irrigation cannot be provided (Knowles 1956). have similar germination temperature requirements, Fairway crested wheatgrass cultivars are best which accounts for the ability of fairway crested adapted to ponderosa pine, mountain brush, pinyon- wheatgrass seedlings to compete favorably with juniper, and benchland sites occupied by big sage- cheatgrass. Asay and Johnson (1983) found that ge- brush. They are also adapted to open, exposed slopes netic differences among populations of crested wheat- with intermixed aspen and spruce-fir communities. grass accounted for over 50 percent of the total pheno- Reitz and others (1936) reported that fairway crested typic variation in traits affecting seeding wheatgrass persists on dry, open areas at elevations establishment. Seed weight was closely correlated to up to 6,800 ft (2,070 m). Plantings in central Utah have seeding establishment. established well on open slopes at elevations near Established stands of fairway crested wheatgrass 8,000 ft (2,440 m). may produce seed crops annually, depending on pre- On adapted sites, fairway crested wheatgrass be- cipitation, but plants do not spread rapidly to adjacent comes dormant during dry periods, but it will resume areas. Seedlings of this species quickly occupy open- growth in fall or spring when adequate moisture ings created in fairway crested wheatgrass stands. becomes available. It tolerates a range of soil textures When seeded in mixtures with other grasses, fairway and pH levels, and withstands saline conditions, but it crested wheatgrass often increases in density at the will not survive flooding for more than a few days. expense of the other species. Walker and others (1995), Fairway crested wheatgrass grows better in moderate reporting on long-term studies of seedings in pinyon- shade than either standard crested wheatgrass or juniper sites in central Utah, found that fairway crested Siberian wheatgrass, but it does not persist in dense wheatgrass density increased at a higher rate than shade. It can establish even with some grazing during native bunchgrasses during periods of normal precipi- seedling establishment (Johnson 1986), although graz- tation, but native bunchgrasses increased in density ing of new seedings is not advised. Mature plants can and ground cover at a greater rate during years of low persist with heavy and repeated grazing. Resistance precipitation. They found that over a 23-year period, to grazing abuse is a primary factor that has promoted native grasses were not able to compete with intro- seedings of this grass. duced species including fairway crested wheatgrass. Fairway crested wheatgrass is generally less well Broadcast seeding fairway crested wheatgrass at low adapted to low-elevation shrubland communities oc- rates of 1 to 2 pounds per acre (1.1 to 2.2 kg per ha) in cupied by Wyoming big sagebrush and salt desert pinyon-juniper and big sagebrush disturbances has species than are other crested wheatgrasses. Fairway been employed to encourage the recovery of native crested wheatgrass plantings have established and species. Although native herbs and shrubs increase persisted in arid regions for a number of years, but initially in some situations, slow but persistent in- some of these have been lost during droughts. The crease of fairway crested wheatgrass followed, and species is suited to sagebrush communities that re- within 25 to 40 years it gained dominance. ceive more than 10 inches (250 mm) of precipitation, but it should not be planted in communities receiving Uses and Management less than this amount. In the Southwest and areas with long warm summers, it will not persist unless Fairway crested wheatgrass has been extensively annual precipitation exceeds 15 inches (380 mm) used in pasture and rangeland plantings in the United (Reynolds and Springfield 1953). States and Canada. In Western Canada, it is used far

314 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Thousands of acres of abandoned farmland and Areas with dense populations of annual weeds should overgrazed rangelands have been seeded to fairway be avoided. Fairway crested wheatgrass plantings crested wheatgrass to provide spring-summer forage have also been important in the control and elimina- for livestock, soil stabilization, and weed control tion of halogeton and Russian thistle (Mathews 1986; (Barnes and others 1952; Bridges 1942; Christ 1934; Mayland 1986). The species has been particularly Hull 1974; Knowles 1956; Westover and Rogler 1947). useful in replacing cheatgrass and slowing the spread In some situations, fairway crested wheatgrass has of wildfires (Monsen and Kitchen 1994). Few other been seeded as a replacement for native grasses (Stark perennial grasses establish as well as crested wheat- and others 1946). Many of these areas were disked or grasses amid weedy competition. burned to remove existing competitive vegetation prior Crested wheatgrass begins growth in early spring to seeding (Malechek 1986). In addition, extensive and provides considerable herbage during this period areas dominated by big sagebrush have been plowed, (Malechek 1986). Nutritive value is good for energy disked, burned, or sprayed to remove the shrubs, then and fair for protein (Dittberner and Olson 1983). seeded with this species and other introduced perenni- Crude protein, calcium, phosphorus, and ash decrease als to enhance livestock grazing (Plummer and others as plants mature in late May and June (Mayland 1955; Robertson and others 1966). Fairway crested 1986). The degree of use is rated as good for livestock wheatgrass seedings have prevented the recovery of and fair for wildlife (Dittberner and Olson 1983). big sagebrush where vigorous stands of the grass have Fairway crested wheatgrass has stabilized eroding developed and the sagebrush seed source has been areas, reduced runoff, and provided a stable vegeta- eliminated. Natural recovery of sagebrush, however, tive cover on many disturbed sites. It performs well as has occurred in other areas. a pioneer species capable of stabilizing exposed sub- If seeded in mixtures, fairway crested wheatgrass strata, and it does not require heavy fertilizer applica- can be drill seeded with other introduced grasses and tions to maintain an acceptable cover. It is frequently broadleaf herbs that produce vigorous seedlings. It is used in big sagebrush communities to control weed capable of persisting with competitive sod-forming invasions and stabilize watersheds, roadways, and perennials, including intermediate wheatgrass and mine disturbances. It furnishes soil protection on sites smooth brome. Although these two sodforming grasses where high-intensity storms are common. may prevent the spread of crested wheatgrass, a In many cases, however, fairway crested wheat- bunchgrass, they do not eliminate it. Fairway crested grass seedings have adversely impacted wildlife habi- wheatgrass is often planted in mixtures with alfalfa in tat. Conversion of native shrub and herb communities mountain brush, ponderosa pine, and other upland to solid stands of this grass have reduced cover and communities to provide summer forage for game and forage for big game and nongame animals (Marlette livestock. Once established, both species persist if and Anderson 1984). Fairway crested wheatgrass properly managed. Production of fairway crested seedings provide important supplements to deer diets wheatgrass may be enhanced by the presence of al- in spring and fall. Elk, pronghorn antelope, and big- falfa. Fairway crested wheatgrass is also frequently horn sheep also use it (Urness l986). These seedings, seeded with alfalfa and late-maturing grasses to main- however, do not afford the diversity and quality of tain herbage quality and diversity (Mayland 1986; forage provided by native communities. A primary USDA Forest Service 1937). concern in seeding fairway crested wheatgrass is the Fairway crested wheatgrass can be broadcast seeded resulting monotypic stands that often develop, reduc- with most revegetation species, including those that ing species diversity and habitat quality (DePuit 1986; are slower to develop if appropriate planting designs Marlette and Anderson 1986). Development of tech- are adopted. Drill seeding fairway crested wheatgrass nology for replacing fairway crested wheatgrass with with slower developing native shrubs and herbs is not more diverse native species to improve habitat charac- recommended unless provisions are made to seed the teristics is becoming a management objective in many species in separate rows or strips that are adequately areas. spaced to minimize competition (Hubbard 1957). Plant- Single species plantings of fairway crested wheat- ing bitterbrush in rows less than 2 ft (0.6 m) from grass normally persist as a monoculture, although fairway crested wheatgrass rows, for example, was invasion of a few other species may occur after 15 to 20 found to significantly decrease vigor and growth of the years. Marlette and Anderson (1984) found that in- shrubs (Hubbard and others 1962). vading species seldom comprise more than 10 percent Fairway crested wheatgrass can be successfully of the total plant density. Stands of fairway crested interseeded in cheatgrass using conventional, deep wheatgrass have persisted for over 50 years in the furrow, rangeland drills that create openings about 4 Great Plains (Mayland 1986), and for over 30 years in to 6 inches (10 to 15 cm) wide. This reduces competi- the Intermountain West (Marlette and Anderson 1984; tion and permits establishment of the perennial grass. Plummer and others 1968). This species does not

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 315 Chapter 18 Grasses invade or migrate into disturbances of early succes- sites receiving at least 12 to 15 inches (300 to 376 mm) sional communities occupied by annual weeds, and it of annual precipitation. Its seedling vigor and ability has made only minor migrations into climax grass- to establish from a 3-inch (7.6-cm) planting depth were lands (Allred 1940). greater than for ‘Nordan’, Fairway, or ‘Ephraim’, and equaled Hycrest (Asay and others 1996). Varieties and Ecotypes Douglas demonstrates promise for seeding in areas dominated by cheatgrass that are prone to frequent Since its introduction in 1906, the crested wheat- burning. It can he used to replace cheatgrass, and it grass complex has been studied intensively. Cultivars can be planted in strips as a fuel or fire barrier. have been developed for a broad range of planting sites Douglas is less drought resistant than standard or and conditions (Asay 1986; Dillman 1946). Selection Hycrest crested wheatgrass, but it remains green and breeding programs are currently underway to longer in summer, and plants are less flammable select strains that are more drought tolerant and during this period. Fires that ignite in cheatgrass may productive, and that remain green longer in summer. burn into the firebreak provided by Douglas, but their Rhizomatous forms of crested wheatgrass are being advance is slowed. developed to provide low maintenance turf and road- The Forest Service, Utah Division of Wildlife Re- way plantings. Breeding has been primarily within sources, Natural Resource Conservation Service, and the diploid or tetraploid levels, but hybrids between the Agriculture Experiment Stations of Utah, Ari- standard crested wheatgrass and induced tetraploid zona, and Idaho released Ephraim crested wheatgrass fairway crested wheatgrass developed by Agricultural in 1983. It was developed from an introduction from Research Service scientists in Logan, UT, are highly Ankara, Turkey, and planted by A. Perry Plummer at competitive on semiarid rangeland sites. an arid range site in central Utah in the early 1940s Scientists at the Agricultural Research Service, Crops where it survived for over 40 years. Seed and plants Research Laboratory in Logan, UT, developed ‘CD-II’ from this site were tested at additional locations crested wheatgrass from the original ‘Hycrest’ hybrid. throughout the Intermountain area (Plummer and Selections were made from Hycrest foundation seed others 1968). fields based on vegetative vigor, absence of purple Ephraim was the first tetraploid cultivar of crested leaves in spring, freedom from disease and insects, wheatgrass released in the United States. It is a and leafiness. From these clonal lines, a second evalu- persistent, sodforming cultivar that produces as much ation was conducted for the same vegetative charac- herbage as Fairway. It is drought tolerant, and can be teristics and for individual seed weight and emergence seeded in areas receiving as little as 10 inches (250 of polycross seed lots from deep seedings. mm) of annual precipitation. Ephraim has persisted CD-II is leafier than Hycrest and produces more when planted in warm, dry areas of southern Utah and forage at low spring temperatures. Stand establish- northern Arizona. It is also well adapted to pinyon- ment and adaptation to arid conditions are compa- juniper and big sagebrush communities of the Inter- rable to Hycrest. Seed production is excellent, ranging mountain region. Plants establishing in moister areas from 600 to 800 pounds per acre (660 to 940 kg per ha). CD-II is recommended for semiarid sites at elevations up to 7,150 ft (2,345 m) receiving from 8 to 18 inches (200 to 450 mm) of annual precipitation (Asay and others 1997). ‘Douglas’ crested wheatgrass, developed by the Ag- ricultural Research Service in Logan, UT, and released in 1994, is the first hexaploid (2n = 42) cultivar of crested wheatgrass released in North America (Asay and others 1995a) (fig. 2). It was named in honor of Dr. Douglas R. Dewey, who initiated research on the perennial Triticeae at the Logan laboratory. Douglas crested wheatgrass was derived from hybrids between a broad-leaved hexaploid accession from the former Soviet Union and hexaploid lines from Iran and Tur- key (Asay and others 1996). Douglas was selected for its leafiness, retention of green leaves into summer, palatability to wildlife and livestock, and high in vitro Figure 2—Mule deer preferred ‘Douglas’ fairway digestibility. It produces larger seeds than diploid or crested wheatgrass to other cultivars of crested wheat- tetraploid cultivars. Douglas should be fall seeded on grass in a fall palatability study (RMRS photo).

316 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses produce rhizomes within 2 to 3 years (Stevens and In 1984 the Agricultural Research Service in coop- Monsen 1985). eration with the Utah Agricultural Experiment Sta- Ephraim is recommended for seeding in pinyon- tion and the Natural Resources Conservation Service juniper, ponderosa pine, mountain brush, and big released Hycrest crested wheatgrass, the first inter- sagebrush communities. It provides excellent ground specific hybrid of crested wheatgrass. Hycrest was cover and protection on disturbed soils. It is highly developed by crossing an induced tetraploid of A. competitive with annual weeds, and can be used with cristatum with natural tetraploids of A. desertorum. Nordan and other cultivars of standard crested wheat- Reciprocal crosses were made to ensure that the cyto- grass to reduce or control cheatgrass, medusahead, plasm of both species would be represented (Asay and and other annual weeds. Seedlings of Ephraim are not others l985). Open pollinated progenies were then as vigorous as those of Hycrest, but once established, established at a nursery site in northern Utah. Eigh- they are extremely persistent in the face of drought, teen clones were selected from these trials and planted cold winters, and heavy grazing. Ephraim furnishes in isolated crossing blocks to provide the first genera- early spring herbage, and greens up in fall if late tion of the synthetic strain (Asay and others 1985). summer or fall moisture is received. It is not shade Seedlings of Hycrest are more competitive than tolerant, but it does persist as an understory in big those of other cultivars. Hycrest competes well with sagebrush and other shrub communities. cheatgrass, halogeton, and other weeds. Seeded Hycrest Ephraim is a competitive cultivar. Established stands stands establish in competition with annual weeds; have not allowed the entry of other perennials, even consequently, this cultivar has been useful in seeding when subjected to heavy and prolonged grazing. sites where other weed-control measures cannot be Ephraim has been successfully seeded on mine distur- fully implemented. In plantings made across the In- bances in northern Nevada. It is also widely used to termountain region, Hycrest has established better plant roadways and other disturbances in the Inter- stands and produced more forage during the first 2 mountain region. It is well adapted to infertile soils years than either Nordan or Fairway (Alderson and and moderately dry conditions. Sharp 1994). Examination of plantings indicates that Fairway crested wheatgrass was the first wheat- seedling vigor, stand establishment, and root growth grass cultivar developed in North America. Released of Hycrest generally exceed those of other crested in 1927, it was selected from P.I. 19536, an introduc- wheatgrass cultivars. Hycrest has proven adapted to tion received from Western Siberia and developed by big sagebrush and pinyon-juniper sites, but it is not as Agriculture Canada (Knowles 1956). It is shorter in drought tolerant as selections of standard or Siberian stature, leafier, more procumbent and winter hardy, crested wheatgrass. but less drought tolerant than most standard crested A diploid cultivar, ‘Parkway’, was developed from wheatgrass cultivars. Although initially developed for Fairway through several generations of recurrent se- open parkways and turf plantings, Fairway has been lection for vigor, forage, and seed production (Alderson widely seeded throughout the Intermountain region, and Sharp 1994). Parkway was released in and has proven better adapted to upland sites than Saskatchewan in 1969 for use in hay and pasture standard crested wheatgrass. It is widely used in the production. Its yields exceed those of Fairway, as the mountain brush, ponderosa pine, pinyon-juniper, and plants are slightly taller, generally 19 to 30 inches (48 mountain big sagebrush zones. Fairway is extremely to 76 cm) in height. This cultivar has not been widely long lived, and has been planted to provide forage, planted on range or wildland sites in the Intermoun- produce ground cover, and prevent the invasion and tain region, but it can be used on many of the same spread of weeds. Fairway seedlings are vigorous and sites as Fairway. It is not recommended for turf capable of establishing amid some competition, but plantings. they are suppressed by dense cheatgrass. Fairway ‘Ruff’ crested wheatgrass is a diploid cultivar jointly reseeds itself well, as an abundance of seeds are released by the Agricultural Research Service and the generally produced, even under semiarid conditions. Nebraska Agricultural Experiment Station in 1974. Fairway is well suited to a variety of site conditions The parental material was derived from Fairway (Asay and is recommended for seeding in mixtures with sod- 1986). Ruff has a spreading bunchgrass habit and forming grasses to provide erosion control. It produces short, leafy culms (Alderson and Sharp 1994). It be- tillers and short rhizomes, and provides excellent gins growth in early spring and is valued as a short- ground cover and high-quality forage. Herbage pro- season pasture plant in drier areas of the Great Plains. duction of Fairway is usually less than that of stan- In the Central Great Plains, its forage production dard crested wheatgrass, but Fairway produces a equals that of Fairway (Alderson and Sharp 1994). Its greater proportion of leaves. With age, standard crested seeds are relatively small, but its seedlings establish wheatgrass develops an abundance of stems that be- well from fall plantings. Ruff competes well with come less palatable to grazing animals. annual and perennial weeds and can be seeded as a

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 317 Chapter 18 Grasses conservation planting to protect soil and control weed invasion in Intermountain areas where Fairway is adapted. Ruff plantings become somewhat sodforming due to the broad bunching habit and seeding within the stand.

Agropyron dasystachyum Thickspike Wheatgrass ______Synonyms Agropyron albicans A. elmeri A. griffthsii A. riparium Elymus lanceolatus

Description Barkworth and Dewey (1985) transferred several species of Agropyron into the genus Elymus, and combined the former taxa A. dasystachyum, A. dasystachyum var. elmeri, and A. dasystachyum var. riparium with E. lanceolatus (Scribn. & Sm.) Gould. Arnow (1987) also combined several Agropyron spe- Figure 3—Thickspike wheatgrass is a widely distrib- cies with E. lanceolatus. Barkworth and others (1983) uted native, sod-forming wheatgrass (photo courtesy and Barkworth and Dewey (1985) described four sub- of Kevin Jensen, USDA ARS, Logan, UT). species of thickspike wheatgrass: E. lanceolatus ssp. albicans (Scribn. & Sm.) Barkw. & D. R. Dewey E. l. ssp. lanceolatus E. l. ssp. psammophilus (Gillett & Senn) A. Love The upper surface of the blade is furrowed, while the E. l. ssp. yukonensis (Scribn & Merr.) Love lower surface is smooth. The upper surface and some- Taxa now included in treatments of E. lanceolatus times the base of the lower surface are roughened. are similar in floral parts and growth form, thus Auricles are about 0.08 inch (2 mm) long. The ligule is simplifying identification. These taxa, however, oc- membranous and up to 0.04 inch (1 mm) long with cupy quite different ecological sites. Montana wheat- ragged edges and a rounded apex. Inflorescences are grass (formerly A. albicans) and streambank wheat- 2.4 to 4.8 inches (6 to 12 cm) long. Spikelets are 0.4 to grass (formerly A. riparium), for example, differ greatly 0.6 inch (1 to1.5 cm) long, each with 3 to 12 loosely to in areas of adaptation and require different manage- closely overlapping florets. Tips of the glumes are ment strategies. subequal and firm, slightly roughened, and often with Thickspike wheatgrass is a native, cool-season pe- small hairs. Glume tips are acute or extended into rennial (Barkworth and Dewey 1985; Hitchcock and short awns. Lemmas are 0.2 to 0.4 inch (6 to 9 mm) others 1969; Looman 1982; USDA Forest Service 1914; long, convex on the back, firm, and five to seven Welsh and others 1987). The root system and long, nerved. They are sparsely to densely hairy, and gener- rapidly developing rhizomes form a sod that varies ally more hairy than the glumes. Lemma tips are with location and subspecies (USDA Forest Service awnless or extended into tiny awns. The palea is about l937; Wambolt 1981) (fig. 3). Plants may develop a as long as the lemma and obtuse at the tip. dense and more or less continuous sod, or they may occur as open, irregular patches intermixed with other Ecological Relationships and Distribution species. Stems are erect, 16 to 32 inches (40 to 80 cm) tall, and most often bluish green. Leaf sheaths have Thickspike wheatgrass is widely distributed, occur- thin, overlapping edges that are smooth to slightly ring in Alaska, southward through the Western United roughened. Blades are 0.04 to 0.12 inch (1 to 3 mm) States and Northern Mexico, eastward across Canada wide, and 2 to 8 inches (5 to 20 cm) long. They are and much of the Great Plains, south to Kansas, and as usually stiff and ascending with slightly rolled edges. far east as West Virginia (Arnow 1987; Ross and

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Hunter 1976). It grows on sites ranging in elevation Fall seeding is recommended. Seeds should be planted from 4,000 to 10,900 ft (1,220 to 3,300 m) that receive about 0.25 to 0.75 inch (0.6 to 1.9 cm) deep by drilling 8 to 20 inches (200 to 500 mm) of annual precipitation. or by broadcasting followed by harrowing. Thick stands Different ecotypes are adapted to soils that are well often develop when thickspike wheatgrass is seeded in drained and sandy, sandy loams, loams, or heavy drill rows on well-prepared seedbeds. When broadcast textured. It occurs on soils that are alkaline or weakly seeded with other species, thickspike wheatgrass tends saline. to form irregular patches that add diversity to wildlife In the Western United States, thickspike wheat- habitat. Development of new stands is aided by rhi- grass is common in sagebrush, pinyon-juniper, moun- zome development. Seedlings of thickspike wheat- tain brush, aspen, moist open parks, and dry meadows grass varieties compete well with annual weeds. The (Wasser 1982). It occurs in desert and chaparral com- species should not be seeded directly with slower munities of the Southwest and in ponderosa pine and developing plants, but it can be seeded in alternate western spruce forests of the Northwest (Arnow 1987; rows or on adjacent sites. Bernard and Brown 1977; Hitchcock and others l969; Sours l983). It is often regarded as a disturbance Uses and Management species, but it is also a major component of grama grass/needlegrass, Idaho fescue, bluebunch wheat- Thickspike wheatgrass is often seeded alone on grass, and mountain brush/aspen communities. disturbances in big sagebrush and mountain brush Thickspike wheatgrass is tolerant of moderate shade communities to stabilize soil and control erosion. It is provided by associated shrubs, but it is not adapted to particularly well adapted to arid environments and dense overstory cover. It recovers quickly following does well on disturbed, slightly alkaline soils. It is fires, and often flourishes in sagebrush and juniper seeded on sites such as mined areas, roadside distur- communities following burning (Wright and Bailey bances, recreation areas, drainageways, and other 1982). It has persisted as a dominant species on sites that receive little maintenance and where a low burned sites for over 30 years (Harniss and Murray sodforming grass is desired. Thickspike wheatgrass is 1973). also commonly used to control the spread of weeds, particularly annual grasses. It is often seeded for this Plant Culture purpose in waste areas and nonirrigated portions of cultivated fields. Thickspike wheatgrass initiates growth in early Thickspike wheatgrass is widely used in range and spring (Dittberner and Olsen 1983). New shoots emerge wildlife plantings. Herbage production is less than for as early as March 1; flowering occurs from May to July. many other grasses, but it does provide excellent Seeds mature in late July and remain on the plant spring-fall and early summer forage. It remains green until mid-August (Blaisdell 1958). Dispersal of ma- later into the summer than cheatgrass. Consequently, ture seeds normally occurs over a 2- to 3-week period. it can be seeded to extend the period of grazing in areas Seed from wildland stands is collected using reel-type infested with this and other annuals. If moisture harvesters mounted on four-wheel-drive vehicles or by becomes available in fall, plants will green up and hand stripping. Seed fields are harvested with conven- provide an important source of forage. Thickspike tional combines. Little debris is collected with the wheatgrass recovers quickly following burning, pro- seeds, as the culms extend above the leaves and the viding herbage and ground cover. It also reduces the seeds shatter readily. Purity of freshly harvested seed spread of cheatgrass and other annual weeds as burned lots usually exceeds 80 to 90 percent with about 85 areas recover. percent germination (Wasser 1982). Most seed used in Populations of thickspike wheatgrass may have re- wildland plantings is from named varieties grown stricted ranges of adaptation. Consequently, attempts under cultivation. Few wildland stands are large should be made to collect and seed local ecotypes. The enough to support extensive harvesting. Seeds are two released cultivars, ‘Critana’ and ‘Sodar’, may not slightly larger than those of crested wheatgrass, aver- be adapted to all sites within the species range. Up- aging about 154,000 per pound (339,500 per kg) land sites, including dry meadows and areas inter- (Wheeler and Hill 1957). mixed with aspen and conifer forests should be planted Seed germination is enhanced by a prechill of about with ecotypes adapted to these areas. 30 days, but some nonstratified seed will germinate in about 20 days under laboratory conditions. Compared Varieties and Ecotypes to other wheatgrasses, germination and seedling development of thickspike wheatgrass is slow, but satis- ‘Bannock’ thickspike wheatgrass, a composite of six factory stands generally develop over time. The seed- collections from Oregon, Idaho, and Washington, was ing rate is sometimes increased to compensate for this released in 1995 by the Natural Resource Conserva- problem. tion Service Plant Materials Center at Aberdeen, ID.

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Bannock is a vigorous, leafy, long-lived, sodproducing develop quickly. Schwendimar is recommended for variety. Its rhizome growth averages about 4 inches areas with neutral, coarse-textured soils that receive 8 (10 cm) per year. It establishes quickly and develops a to 20 inches (200 to 500 mm) of annual precipitation. protective sod on dry sites. It is a more effective The variety can be used to quickly stabilize well-drained, sodproducer than Sodar when seeded in the 8- to 10-inch droughty sites. It can also be seeded in mixtures to (200- to 250-mm) precipitation zone. Plants are 18 to provide mid to late summer forage for wildlife and 24 inches (45 to 60 cm) tall, and may grow to 40 inches livestock. Plantings have persisted on a wide range of (100 cm) in height when irrigated. Leaves are abundant sites in the Columbia River Basin, but the variety has and are produced 8 to 12 inches (20 to 30 cm) up the stem. not seen extensive use outside this area (USDA Soil Bannock is recommended for seeding in mixtures on Conservation Service 1994). arid rangeland sites and for conservation plantings to Sodar was released as a streambank wheatgrass provide soil protection. It is useful for revegetating variety, but it is now considered a thickspike wheat- mined lands and other disturbances, and its highly grass. The Idaho and Washington Agricultural Ex- competitive sod reduces the spread of cheatgrass and periment Stations cooperatively released it in 1954. other annual weeds (USDA Natural Resourcess Con- Sodar was developed by several generations of mass servation Service 2001). Inflorescences are large and selection from material collected near Canyon City, the florets are awnless. Seed yields range from 250 to Grant County, OR, and grown at the Natural Resource 400 pounds per acre (275 to 440 kg per ha). Conservation Service Plant Materials Center, Aber- The Natural Resource Conservation Service Plant deen, ID (Alderson and Sharp 1994). Materials Center at Bridger, MT, and the Montana Sodar is a rhizomatous, drought-resistant peren- and Wyoming Agriculture Experiment Stations coop- nial, 20 to 30 inches (5 to 7.6 dm) tall that is used eratively released Critana thickspike wheatgrass in primarily for erosion control and conservation 1971. D. E. Ryerson collected the original material plantings. It is more drought tolerant and widely from a roadcut near Havre, MT, in 1960. Critana adapted to big sagebrush communities than Critana. provides a low-stature ground cover and a dense sod In the Intermountain region, Sodar is seeded at eleva- that requires little maintenance. This cultivar exhib- tions below 3,200 ft (975 m) on sites that receive 10 to its excellent seedling vigor and establishes well, even 12 inches (250 to 300 mm) of annual precipitation. in areas receiving low rainfall. Plants are strongly Sodar receives use on roadways, dry drainageways, rhizomatous and produce low to moderate amounts of destabilized watersheds, and mine disturbances. Sodar herbage. Leaves are small, fine, and palatable. Plants seedlings are vigorous and compete well with annual are tolerant of grazing and produce useful spring and weeds; established stands restrict weed spread. early summer forage. Critana is best adapted to the Sodar is not commonly seeded to provide livestock Pacific Northwest, the Intermountain West, and the forage because of its low herbage production. Plants Western Great Plains. It is commonly seeded in Mon- begin growth as early as standard crested wheatgrass tana, Wyoming, and the western Dakotas on range- and provide good spring forage. They remain green lands and mine sites receiving 10 to 16 inches (250 to somewhat later in summer than standard crested 400 mm) of annual precipitation (Thornburg 1982). wheatgrass. The presence of green leaves into the dry Critana can be used to stabilize infertile disturbances, season can reduce the spread of fire. Sodar is more provide soil protection in areas where some traffic can drought tolerant than most populations of bluebunch be expected, and control weeds. It has been success- wheatgrass. Fall regrowth is generally limited. Sodar fully seeded and maintained along roadways, airport provides diversity and ground cover in dry communi- runways, dry drainageways, recreational areas, and ties where few other species are adapted. It can be mine disturbances (Alderson and Sharp 1994). established easily by drilling or by broadcast seeding ‘Schwendimar’ thickspike wheatgrass was released and harrowing. in 1994 by the Natural Resources Conservation Service Plant Materials Center at Pullman, WA. It was named for John Schwendiman who collected seed from its site Agropyron desertorum of origin, a sandy, windblown area on the banks of the Standard Crested Wheatgrass or Columbia River east of the Dallas, OR, in 1934. Desert Crested Wheatgrass ______Schwendimar is a rhizomatous, cool-season perennial with bluish, glaucous culms. It develops a dense and Description fibrous but shallow root system. Some roots extend more than 2 ft (0.6 m) from the plant. Seeds are Standard crested wheatgrass is an introduced, cool- relatively large. Seedlings establish well from spring or season, generally nonrhizomatous, perennial bunch- fall plantings. They are no more vigorous than those of grass that is similar to fairway crested wheatgrass in other thickspike wheatgrass cultivars, but stands appearance (figs. 1 and 4). Dewey (l986) described

320 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses characteristics that can sometimes be used to differ- Arnow (1987) combined A. cristatum, A. desertorum, entiate the two species. The spikelets of standard and A. fragile (= A. sibiricum) into a single species, A. crested wheatgrass are subcylindrical and oblong to cristatum, based on their morphological similarities. linear, diverging from the rachis at angles of 30 to 45 She also provided a key for separating standard and degrees. The glumes of standard crested wheat- fairway crested wheatgrasses in the field using many grass are appressed to the lemmas, and the lemmas of the same floral characteristics described by Dewey support a short, straight awn that is 0.1 inch (3 mm) (1986). This key described the culms of fairway crested or less in length. Fairway crested wheatgrass has wheatgrass as often less than 1.3 ft (4 dm) tall and broad, pectinate spikes and spikelets that diverge typically variable in height with both basal and cauline from the rachis at much greater angles (45 to 90 leaves. Culms of standard crested wheatgrass were degrees), and its glumes are not appressed to the described as often more than 1.3 ft (4 dm) tall and lemmas, thus the spike has a bristly appearance. Its typically more or less equal in height with strictly lemmas are short with straight awns that may reach basal leaves. 0.2 inch (5 mm) in length. Spikes of standard crested wheatgrass are 6 inches (15 cm) long and 0.2 to 0.9 inch (5 to 23 mm) wide. Spikelets are 0.2 to 0.6 inch (5 to 15 mm) long, excluding the awns, sessile, and spaced 0.04 to 0.12 inch (1 to 3 mm) apart along the rachis. Each spikelet produces 3 to 10 florets, but the upper florets are sterile. Glumes are nearly equal, about 0.28 inch (7 mm) long, and linear to lanceolate with the tip often tapering to a short awn. Glumes and paleas are often rough along the keel. The lemma is 0.16 to 0.33 inch (4 to 8.5 mm) long and sometimes short pubescent. Leaf blades are 2 to 8 inches (5 to 20 cm) long and 0.08 to 0.2 inch (2 to 5 mm) wide. Most are flat or slightly bent along the midrib. Plants reproduce by seeds and tillers (Arnow 1987; Hitchcock 1950).

Ecological Relationships and Distribution Standard crested wheatgrass is native to European Russia, the Caucasus Mountains, Western Siberia, and Central Asia (Weintraub 1953). It has been widely planted throughout North America in dryland and pasture seedings. In the Western United States it is extensively seeded in salt desert shrublands and in big sagebrush, pinyon-juniper, and mountain big sagebrush communities (Johnson 1986). Standard crested wheatgrass is typically less tolerant of shade, high- elevation conditions, and salty soils than fairway crested wheatgrass (Plummer and others 1958). Standard crested wheatgrass is well adapted to the Northern Great Plains (Westover and others 1932) and the prairies of Western Canada (Hubbard 1949). Sarvis (1941) reported that crested wheatgrasses, including standard crested wheatgrass, progressed from a promising species in 1917 to the leading hay and pasture species in the Northern Great Plains. Lorenz (1986) reported that it had replaced smooth brome as the most important species planted in the Northern Great Plains of the United States and in Figure 4—Standard crested wheatgrass is a Eur- Canada. It exceeds smooth brome in drought toler- asian introduction that has been widely seeded in ance, forage production, seed production, ease of es- semiarid areas of the Western United States tablishment, and preference by livestock as hay or (RMRS photo). pasture.

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Standard crested wheatgrass has been seeded widely lands averages about 250 to 300 pounds per acre and has proven well adapted in almost all big sage- (280 to 336 kg per ha) (Mayland 1986). For cultivated brush communities in the West. It is recognized for its fields in the Great Plains, Intermountain region, and ability to establish and persist in disturbances on the Southwest, yields vary between 100 and 600 semiarid lands. It is adapted to the high desert of pounds per acre (112 to 672 kg per ha) (Reynolds and eastern Oregon and similar areas throughout the Springfield 1953; Westover and others 1932). Wide Pacific Northwest (Jackman and others 1936). row spacings, 24 to 36 inches (61 to 91 cm), increase Leckenby and Toweill (1983) reported that it per- yields (Windle and others 1966). Knowles and Kilcher formed well in juniper/big sagebrush-antelope bitter- (1983) reported maximum yields resulted when stands brush communities in Oregon. It is well adapted to the were planted at 12 to 18 inches (30 to 46 cm) row cold desert regions of Utah and Nevada. Standard spacings and fertilized with about 40 pounds of nitro- crested wheatgrass and Russian wildrye have per- gen per acre (45 kg per ha). Narrow spacings of less sisted better than any other species tested in seedings than 18 inches (46 cm) are not advisable. McGinnies on winterfat/shadscale sites in south-central Utah. (1971) found that during periods of below average These two introduced grasses demonstrate the ability rainfall, plantings with row spacings of 18 to 24 inches to persist for over 60 years without significant loss of (46 to 60 cm) failed to produce harvestable crops. vigor or stand density. Standard crested wheatgrass is Seed production continues for many years on fields also used in the warm desert regions of the Southwest, that are kept free of weeds. Seeds shatter when ma- particularly pinyon-juniper and big sagebrush com- ture, and they must be harvested before being dis- munities of northern New Mexico and Arizona lodged by wind. Some debris can be removed during (Reynolds and Springfield 1953) and the mid and the combining process. Once seeds are harvested, they lower elevation plant communities of the Four Cor- are easily threshed and cleaned using fanning mills. ners area. Seed quality standards are normally 95 percent for Although it is widely planted, standard crested purity and 90 to 95 percent for germination. Seeds can wheatgrass it is not suited for some situations where be stored for 2 to 3 years or longer without appreciable it has been seeded (Hull 1963b; Johnson 1986). Stan- loss in germination or seedling vigor. dard crested wheatgrass is adapted to areas receiving Seeds germinate rapidly when placed in cool, moist 10 to 20 inches (250 to 500 mm) of annual precipita- environments. Love and Hanson (1932) found that tion. Stands often establish and persist on sites receiv- seeds of crested wheatgrass placed in incubation at 25 ing less than 10 inches (250 mm) of annual precipita- ∞C (77 ∞F) produce a coleorhiza and several root hairs tion, but favorable conditions must exist at the time of within 24 to 48 hours. Both the coleoptile and primary seeding and seedling emergence to assure establish- root were well developed within 72 to 96 hours. This ment. Seedings have persisted for many years in very explains, in part, why crested wheatgrass plants es- dry situations, but eventually succumb during periods tablish so well under arid conditions. Seedlings are of extreme drought. Standard crested wheatgrass is capable of emerging and establishing within a short better adapted to dry sites than varieties of fairway period before soil surfaces dry in spring. Plummer crested wheatgrass, but it is less salt tolerant (Maas (1943) and Plummer and others (1958) concluded that and Hoffman 1977). Dewey (1960) observed differ- the widespread success of crested wheatgrass estab- ences in salt tolerance among collections of both spe- lishment is due to the greater root length attained by cies, but concluded Fairway crested wheatgrass gen- developing seedlings at low temperatures when com- erally produced greater yields when grown under salt pared to many natives. stress than did standard crested wheatgrass. Seedlings can be established from both fall and spring plantings, but fall seedings are recommended. Plant Culture Seeds are drill seeded or broadcast and covered using a chain or harrow. Standard crested wheatgrass is Standard crested wheatgrass generally produces often seeded on rough soil surfaces without any at- good seed yields when grown under cultivation on tempt to cover the seed. Although this practice is not irrigated or nonirrigated sites. It is highly competitive advised, satisfactory stands are sometimes achieved. and can normally be grown without serious weed Planting at depths of about 0.5 to 0.75 inch (1.3 to 1.9 problems. Seed yields vary among years, but some cm) enhances seedling emergence. Deep-furrow drill- seed is normally produced. Considerable seed is har- ing has been recognized as a satisfactory means of vested from dry farms or wildland stands during years planting rangeland sites (McGinnies 1959), especially of above average precipitation. Stable markets have where weeds are a factor. Evans and others (1970) developed for this species, and seed prices are gener- concluded that deep furrows provide a favorable seed- ally low. bed by reducing temperature extremes and moisture Seed production fields can be established rather loss. quickly, normally within 2 years. Seed yield on dry

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Hull and Klomp (1967) reported that drilling pro- this perennial wheatgrass are not likely to revert to a duces 10 times more seedlings than broadcasting and native composition. that drilled stands reach maturity much sooner. This is important if a single species is desired, but it Uses and Management produces more competition with other species, including recovering natives. Seeding closely spaced furrows Standard crested wheatgrass has been used for a at high rates is advisable for weed control. Broadcast number of purposes. Westover and Rogler (1947) con- seeding followed by chaining or harrowing to cover cluded that this grass has considerable value as a the soil can produce successful plantings. Broad- supplement to native range. It is perhaps the most cast plantings are not as prone to frost damage as widely seeded grass in the Intermountain region, drill seedings that promote more uniform seedling because it is adapted to semiarid sites where native emergence. communities have been seriously altered, and where Mixed plantings of standard crested wheatgrass few other species establish. It can be grazed 2 to 3 and less competitive, slower developing species can be weeks earlier than most native ranges, providing established by interseeding the species in alternate highly succulent forage for lactating livestock in early rows if adequate row spacing is provided (McKell spring (Harris and others 1968). Considerable re- 1986). Hubbard and others (1962), for example, found growth becomes available if fall precipitation occurs. that antelope bitterbrush must be seeded at least 2 ft The ease with which this species establishes and its (0.6 m) from crested wheatgrass to assure initial early spring growth favor its use to provide soil protec- seedling survival. Walker and others (1995) found tion and control weeds such as halogeton and cheatgrass that mixed seedings of standard crested wheatgrass (Sharp 1986; Stoddart and others 1975). It is particu- and other introduced and native species on pinyon- larly important for seeding semiarid and arid sites juniper sites declined in diversity over a 30-year pe- (Lorenz 1986). Standard crested wheatgrass gained riod. Crested wheatgrass increased at the expense of early recognition for its use in restoring degraded native plants during average or above average precipi- ranges during the drought of the 1930s (Lorenz 1986; tation years. Monsen and Shaw (1983c) reported that Sarvis 1941). It remains an important species for crested wheatgrass, seeded as an understory with seeding abandoned and degraded rangelands and is antelope bitterbrush, restricted shrub seedling re- considered one of the most important species for weed cruitment over a 40-year period. Standard crested and fire control in big sagebrush and salt desert wheatgrass reduces natural reestablishment of big communities. sagebrush and associated species in areas where an- Standard crested wheatgrass has also been used to nual precipitation is near the lower limit for the native convert big sagebrush and native bunchgrass ranges species. to crested wheatgrass monocultures. Extensive sage- Recommended seeding rates vary considerably brush acreages have been cleared and seeded with among regions. Rates of 8 to 12 pounds per acre (9 to standard crested wheatgrass to provide a more palat- 13.5 kg per ha) were commonly employed in early able species for livestock forage (Frischknecht 1963). plantings. Drill seeding rows 6 to 12 inches (15 to 30 Some managers have promoted such conversions of cm) apart at this rate provides competitive cover that native bunchgrass ranges because standard crested will prevent weed invasion. For other plantings we wheatgrass is more tolerant of grazing. Standard recommend rates of 1 to 2 pounds per acre (1.1 to 2.2 crested wheatgrass sometimes provides spring forage kg per ha) to permit initial establishment of other 2 or 3 weeks earlier than some native grass and brome seeded species and recovery of existing native herbs grass pastures (Williams and Post 1941), but not as and shrubs. Attempting to manage for natural recov- early as Sandberg bluegrass or muttongrass. Com- ery by seeding standard crested wheatgrass with na- pared to bluebunch wheatgrass, standard crested tive species is, however, a questionable practice, as full wheatgrass can more rapidly activate buds, initiate stands of the wheatgrass may develop over time. new tillers, and better allocate limited carbon reserves Standard crested wheatgrass does not spread into to support regrowth (Caldwell and Richards 1986b). adjacent areas, but seedling recruitment does occur Forage quality of standard crested wheatgrass is within the planting. Few seedlings of other species high during May and June, but declines rapidly as establish in dense wheatgrass stands because the flowering begins (Malechek 1986). For this reason it is existing crested wheatgrass plants and emerging seed- often seeded with perennial broadleaf herbs, includ- lings are quite competitive (Marlette and Anderson ing alfalfa, to enhance and sustain high-quality herb- 1986). As large disturbances are seeded to standard age later in the growing season. Mixed grass-legume crested wheatgrass, potential native seed banks are seedings also substantially increase yields (Johnson depleted over time, and recruitment of new plants is and Nichols 1969; Schuman and others1982) and are limited to the edges of the seedings. Areas seeded to recommended for sites exceeding 14 inches (350 mm) of annual precipitation.

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Mixed seedings must be carefully managed to en- fields. To be fully successful, however, revegetation sure that all species become established and persist. efforts must allow for the recovery of native species, Competition from the grass coupled with selective and standard crested wheatgrass prevents this from grazing of legumes can restrict natural reseeding of occurring. Restoration of diverse native plant commu- the legumes. Rosenstock and Stevens (1989) found no nities is essential to maintain ecologically sound and recruitment of alfalfa in mixed crested wheatgrass properly functioning systems. Converting extensive seedings on pinyon-juniper sites in central Utah and areas to a single introduced species is not advised. attributed this to competition from established grasses. Standard crested wheatgrass is a persistent peren- Mixed seedings of standard crested wheatgrass and nial that can withstand annual grazing. Reynolds and forage kochia have been planted to provide midsum- Springfield (1953) recommend that use not exceed 55 mer and winter grazing not provided by the grass percent in any year. Many stands seeded between alone (Monsen and others 1990). Cattle graze these 1932 and 1934 in southeastern Idaho have been grazed seedings during periods when the grass is dry and not each year, and persist after 60 years (Hull and Klomp grazed. 1966). Standard crested wheatgrass has been used as Standard crested wheatgrass has been widely used a hay crop in dryland pastures (Stitt 1958), but more to seed drastic disturbances including mined lands, productive and better quality herbage is produced roadways, and degraded watersheds. It establishes from mixed plantings. This species can be used as and persists well on such sites and provides weed winter forage if supplements are provided. control and soil protection (DePuit 1986). It can be damaged by wildfires, but it normally recovers unless Varieties and Ecotypes the burn occurs under extreme conditions. The decision to use standard crested wheatgrass in Nordan, released in 1953, is the most widely used wildlife improvement projects must be carefully con- standard crested wheatgrass variety in the United sidered. Because it is a competitive species that fre- States. It was selected from material received from the quently gains dominance (Johnson and Nichols 1969; U.S.S.R. and planted at Dickerson, ND, by the Agri- Schuman and others 1982; Stitt 1958), important cultural Research Service, Northern Great Plains habitat can eventually be lost. Increases in crested Research Center. Nordan was selected for its large wheatgrass can also occur over periods of 10 to 40 seeds, reduced awns, and good seedling vigor (Rogler years or more without livestock grazing (Monsen and 1954; Wolfe and Morrison 1957). It is a uniform, erect, Anderson 1993). Introduced grasses, including both and productive cultivar with excellent establishment standard and fairway crested wheatgrass, have been attributes. Plants are palatable when green, but pal- used to seed degraded big sagebrush, salt desert shrub, atability is reduced as the dry stems attain maturity. and pinyon-juniper communities. In some situations, Nordan generally produces high amounts of seed each these species are used to replace natives. Although year, even under arid conditions. This cultivar is both grasses furnish early spring herbage, they are not widely adapted to sagebrush steppe communities, but the species most preferred by mule deer (Austin and it is also seeded in salt desert shrublands. It is one of others 1994) or pronghorn antelope (Urness 1986; the main cultivars planted in large restoration projects Yoakum 1980). Memmott (1995) reported that the on semiarid lands. nutritive qualities of various native shrubs exceeded ‘Summit’ was developed and released in 1953 by those of crested wheatgrass in nearly all months. Agriculture Canada at Saskatoon, Saskatchewan Seeding standard crested wheatgrass and other highly (Hanson 1959). This cultivar is similar to standard competitive introduced grasses has reduced the pro- crested wheatgrass varieties grown in the United ductivity of sage-grouse habitat. These grasses have States. It was developed from materials obtained from greatly decreased the density and distribution of forbs the Western Siberian Experiment Station. Fairway- and sagebrush in seeded communities and inhibited type plants were removed during the first year of their natural establishment and successful reproduc- increase (Alderson and Sharp 1994). Summit provides tion. Yoakum (1983) concluded that broadleaf herbs abundant forage, but problems associated with seed and woody species are more important to pronghorn processing have limited its popularity and use (Asay antelope than are seeded grasses. and Knowles 1985b). Elk herds have benefited immensely from large The Agricultural Research Service in Logan, UT, is revegetation projects in Utah. Animals have been currently investigating other selections of crested attracted to these areas for winter grazing, and they wheatgrass. Attention is being given to the develop- use grass-herb seedings at all seasons. Sites seeded to ment of a turf-type, more decumbent, and rhizoma- standard crested wheatgrass, alfalfa, small burnet, tous cultivar. In addition, studies are being conducted and other grasses have successfully attracted animals to combine leafiness, retention of forage quality, larger in early spring, thus reducing damage to agricultural seed size, and vegetative vigor with the establishment traits and early growth of Hycrest.

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Agropyron elongatum but discarded as being too aggressive. The species received favorable ratings in Athens, GA, in 1920, but Tall Wheatgrass ______it was ignored for another decade (Weintraub 1953). Synonyms By 1930 newer introductions and studies prompted increased use of tall wheatgrass in the Western United Agropyron elongatum ssp. ruthenicum States. Plants discovered growing in salt marshes Elytrigia ponticum were subsequently found well adapted to saline or Elymus elongatus ssp. ponticus alkaline soils of the Intermountain region. Tall wheat- Lophopyrum ponticum grass is adapted to subirrigated and irrigated saline soils and poorly drained alkali soils at elevations to Description 6,560 ft (2,000 m) (Hafenrichter and others 1968). Seeding tall wheatgrass has increased forage produc- Tall wheatgrass is an introduced, perennial, cool- tion and quality on greasewood and saltgrass sites season, bunchgrass (fig. 5). It is a tall, very coarse, receiving 8 to 12 inches (200 to 300 mm) of annual productive, and late-maturing species. Culms are precipitation. Tall wheatgrass is also adapted to sage- densely tufted, erect, leafy, and 16 to 42 inches (4 to 10 brush and mountain brush rangelands, but it has not dm) tall. Leaves are flat or involute, 0.08 to 0.3 inch (2 performed well on acid soils of the Idaho Batholith. It to 8 mm) wide, and glabrous or with the upper surface survives extremely cold winter temperatures, even scabrous or pubescent with short stiff hairs. Spikes when soils are bare of snow. It is widely used in the are generally erect and 3 to 16 inches (8 to 40 cm) long Intermountain region, the Great Plains, and the North- with a continuous rachis. Internodes are mostly 0.3 to western United States, and parts of South America. 0.8 inch (7 to 20 mm) long. Spikelets are solitary at each node, compressed at maturity, and 0.5 to 1.4 Plant Culture inches (13 to 35 mm) long. Each produces five to eight flowers. Glumes are typically 0.2 to 0.4 inch (6 to 11 Seed of tall wheatgrass is produced in cultivated mm) long, thick, hardened at maturity, conspicuously fields at mid- and low-elevation locations character- five to seven nerved, and rounded on the back. Lem- ized by long growing seasons (Hafenrichter and others mas are 0.4 to 0.5 inch (9 to 13 mm) long, thick, 1968). Seed fields should have deep, well-drained hardened, oblong to lanceolate, and truncate to acute soils. Recommended row spacing is 36 to 42 inches (90 at the apex (Arnow 1987). to 106 cm). Fields must be cultivated to control weeds and maintain proper spacing. Plants are leafy and Ecological Relationships and Distribution quite tall, sometimes exceeding 6.5 ft (2.0 m) in height, but they must be irrigated and fertilized with nitrogen Tall wheatgrass is a native of southern Europe and at 60 to 80 pounds per acre (67 to 89 kg per ha) to Asia Minor, where it occurs on saline meadows and maximize yields. Seeds ripen in late summer, particu- seashores (Beetle 1955). Selections from Russia and larly when grown under irrigation. Foliage and dry Turkey were introduced in the United States in 1909, seeds can be harvested with a combine when dry. If plants are too green to harvest directly, stands can first be cut, windrowed, and allowed to dry before combining. Yields vary from 300 to 500 pounds per acre (333 to 555 kg per ha) (Hafenrichter and others 1968). If properly managed, seed fields will remain productive for many years. Field-harvested materials are easily cleaned, as the seeds are large, smooth, and easily detached from the spikelet. Seed lots with over 90 percent purity and germination are commonly sold. Late fall or early winter seeding is recommended for establishing tall wheatgrass in upland rehabilitation plantings. Meadows and wetlands that do not dry out in early spring may be seeded in late spring and summer. Sites requiring control of excess salinity or weedy competition may also be seeded in these seasons. Surface salts can be flushed from the seedbed by spring and early summer irrigation; mechanical weed Figure 5—Tall wheatgrass was introduced from control treatments may be continued until midsum- southern Europe and Central Asia and is adapted to mer. Spring and summer seeded sites usually require saline and alkaline soils (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 325 Chapter 18 Grasses irrigation before and after planting to assure seed the summer. As plants mature, the leaves become germination and seedling establishment. coarse and less palatable, but even in winter large Seeds must be incorporated into the soil at 0.5- to amounts are consumed by livestock and game ani- 1.0-inch (1.2- to 2.5-cm) depths; deep furrow drilling is mals. Heinrichs and Carson (1956) found that the recommended. If surface salts are a problem for small protein content of tall wheatgrass was lower and the seedlings, the deep furrows provide ridges where evapo- crude fiber content higher at all seasons than for other ration can occur, leaving salts on the ridge surfaces grasses used in dryland revegetation. Cook and others and reducing the salt content in the bottom of the (1967) also reported lower levels of total protein for tall furrows where the seeds are planted. wheatgrass compared to selected species planted in Tall wheatgrass has large seeds that can be planted central Utah. Total digestible nutrients and digestible with most equipment. Seeds germinate rapidly. protein for tall wheatgrass, however, are higher in the Germinants are vigorous and develop quickly, produc- early leaf stages than for crested wheatgrass ing seedlings that compete well with weeds. Tall (Hafenrichter and others 1968). wheatgrass is often seeded alone, but it can be success- Tall wheatgrass can be seeded in irrigated and fully established in mixed plantings. nonirrigated pastures. On rangelands, plants may reach heights of more than 3 ft (1 m), while irrigated Uses and Management plants often attain heights of nearly 6 ft (1.8 m). Although tall and rank, tall wheatgrass produces Tall wheatgrass is adapted to valley bottoms that abundant forage. It is often used for hay and provides may be saline or alkaline, and to foothill communities. good silage when cut between the heading and soft- It is the most robust and erect bunchgrass currently dough stages. On rangelands it responds well to fall under use for range and pasture plantings. On salty rains and greens up as moisture becomes available. soils it has established better and produced higher With irrigation, plants provide green forage through- yields than most other forage species tested. When out the summer. It does not recover as well from seeded on low-value saltgrass sites it produces as clipping as other pasture plants, but it can be grazed much as 7 tons of forage per acre (15.5 metric tons per during all seasons. Tall wheatgrass is often seeded ha) (Richards and Hawk 1945) and 600 pounds of seed alone and grazed during midsummer. Sheep as well as per acre (660 kg per ha). It will grow on sites with high cattle use this species. Young lambs and ewes forage water tables, but it will not survive flooding for ex- on tall wheatgrass from early spring to mid to late tended periods. summer. Horses and dry cattle are often wintered on Dewey (1960) found that tall wheatgrass was supe- tall wheatgrass pastures. rior to other wheatgrass species in its ability to grow Tall wheatgrass is generally seeded with other spe- on salt affected soils. Salinity levels of 6,000 to 18,000 cies on salt desert shrublands and in sagebrush and ppm increase yields of this species, resulting, in part, pinyon-juniper communities. Its production may equal through reduction in competition with other plants. or exceed that of crested wheatgrass under these Forsberg (1953) found tall wheatgrass and slender conditions. It generally develops a dense stand when wheatgrass were more resistant to saline conditions seeded on seasonally dry meadows and lowlands, but than other plants tested. These two species persisted it forms more open stands on foothill benchlands. Once in soils with conductivity readings as high as 15.13 established, plants persist well, but spread from natu- mmhos per cm. Roundy and others (1983) found that ral seeding is quite slow. tall wheatgrass and Great Basin wildrye cultivars Tall wheatgrass is useful for planting disturbed waste- were well suited to degraded saline and alkaline rangelands where soil salinity prevents the establishment or lands in arid regions of the West. Both species sur- growth of other species. It is planted along waterways vived soil osmotic potentials as low as –3.5 MPa but and on roadway disturbances, abandoned croplands, grew very little below –1 MPa. Robertson (1955) re- and sites where grazing has eliminated native species. ported that roots of tall wheatgrass penetrated to It is useful for controlling annual weeds, particularly depths of more than 12 ft (3.7 m) in saline or alkaline chenopods. Tall wheatgrass is a useful cover crop and soils. can be used to provide nesting cover, winter protection, Tall wheatgrass is less drought tolerant than stan- and escape cover for upland game birds. It is commonly dard crested wheatgrass, but it is considerably more used to protect the soil and provide habitat for wildlife drought tolerant than smooth brome. It does well on on farmlands where it is adapted. rangelands that receive more than 12 inches (300 mm) of annual precipitation. It provides excellent spring Varieties and Ecotypes forage, as it is about 2 weeks earlier than most native vegetation. It is the latest maturing grass adapted to ‘Alkar’ tall wheatgrass was released in 1951 by the the continental climates of the West (Hafenrichter National Resources Conservation Service Plant Mate- and others 1968). Some leaves remain green most of rials Center, in Pullman, WA. It was developed by

326 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses mass selection from spaced plantings of materials Description introduced from the U.S.S.R. in 1932 (Hanson 1959). Alkar is a tall, very late maturing bunchgrass with Intermediate wheatgrass is an introduced, cool- large, coarse, blue-green leaves. This cultivar pro- season, sodforming perennial (fig. 6). Stems are bluish duces large seeds, and its seedlings are vigorous and green, robust, erect, and 16 to 48 inches (40 to 120 cm) fast growing. Alkar is tolerant of wet, alkaline soils tall. Leaf sheaths are typically ciliate, at least along and can be seeded on semiarid rangelands at eleva- one margin. Blades are 4 to 16 inches (10 to 40 cm) tions between 290 and 5,500 ft (90 to1,830 m). Because long, 0.2 to 0.4 inch (5 to 10 mm) wide, dark green to of its palatability, it receives use as a pasture plant on bluish green, and flat or involute. Blade surfaces are subirrigated and irrigated saline and alkaline soils distinctly veined and glabrous to pubescent with short, (Alderson and Sharp 1994). stiff hairs. The ligule is generally flat, 0.4 to 0.8 inch ‘Jose’ was released in 1965 by the National Re- (1 to 2 mm) long, membranous, and smooth to ragged sources Conservation Service Plant Materials Center along the edges. Auricles are well developed. Spikes at Los Lunas, NM. It was selected from material are erect, 4 to 8 inches (10 to 20 cm) long, and slender. received from Eurasia. Jose was developed as a pas- Internodes of the rachis are 0.24 to 0.80 inch (6 to 20 ture and hay crop. It is a uniform, medium-tall bunch- mm) long near the center of the spike. Spikelets are grass with green leaves. It is shorter and produces less solitary at each node, compressed at maturity, and seed and herbage than other tall wheatgrass cultivars. overlap only slightly, if at all. They are 0.35 to 0.8 inch It is drought tolerant and very salt tolerant (Alderson (9 to 20 mm) long, 0.8 to 0.12 inch (2 to 3 mm) wide, and and Sharp 1994). three to eight flowered. Glumes are thick and hard- ‘Largo’ tall wheatgrass was cooperatively released ened, oblong to lanceolate, conspicuously three to in 1961 by the National Resources Conservation Ser- seven nerved, and rounded to slightly keeled on the vice Plant Materials Center in Los Lunas, NM, the back with a blunt or pointed tip. Spikelets are awn- Utah Agricultural Experiment Station in Logan, UT, less or awn tipped (Arnow 1987; Hitchcock 1950; the Agricultural Research Service, Logan, UT, and the Stubbendieck and others 1992). New Mexico Agricultural Experiment Station, Las Pubescent wheatgrass was previously considered a Cruces, NM. It was developed as a forage species for separate species from intermediate wheatgrass (Nevski seedings in Colorado, Utah, Arizona, and New Mexico, 1934), but they differ only in the presence or absence but Jose has largely replaced it. of a pubescence on the spikes, seeds, and leaves. ‘Orbit’ was selected at the Agriculture Canada Re- search Station, Swift Current, Saskatchewan, in 1966. Alkar was selected for its winter hardiness, seed yields, and forage production. This cultivar is superior to Alkar in winter hardiness, but similar in seed and forage yields. It can withstand spring flooding for 3 to 4 weeks (Alderson and Sharp 1994). ‘Platte’ was selected at the Nebraska Agriculture Experiment Station in Lincoln, NE, and released in 1972 by the Agricultural Research Service and the University of Nebraska, Lincoln. It is a winter-hardy bunchgrass that spreads from short rhizomes and produces long, narrow leaves and rather large seeds. It is primarily used to seed saline and alkaline soils in the Great Plains (Alderson and Sharp 1994).

Agropyron intermedium Intermediate Wheatgrass ______Synonyms Agropyron trichophorum Elymus hispidus Elytrigia intermedia Figure 6—Intermediate wheatgrass, a sodforming Thinopyrum intermedium introduction, is seeded to provide hay, forage, and Thinopyrum intermedium ssp. barbulatum stabilization of disturbed sites (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 327 Chapter 18 Grasses

Dewey (1977) concluded that these are not sufficient Plant Culture differences for separation of species within this tribe, particularly as the taxa do not differ in greatly in Intermediate wheatgrass seed is produced in geographical distribution, ecological adaptation, mode nonirrigated and irrigated fields. Yields range from of reproduction, or cytological characteristics. Consid- 250 to 500 pounds per acre (280 to 560 kg per ha) erable variability, however, does occur within each (Hafenrichter and others 1968). Good yields can be form, and cultivars of each have been developed and produced with limited fertilization and irrigation. released. Plants develop a competitive sod rather quickly, and renovation of seed production fields with mechanical Ecological Relationships and Distribution tillage is required to maintain acceptable seed production. Intermediate wheatgrass produces seedstalks Intermediate wheatgrass and its pubescent form, with large seedheads. Seeds ripen uniformly and re- pubescent wheatgrass, are distributed from southern main on the plant, but can be easily separated from the Europe through the Middle East to Western Pakistan inflorescence during cleaning. Immature florets can (Bor 1970). Both forms of the species were introduced be removed using fanning mills and other separators. into the United States from Central Asia in 1907 (Asay Seeds can be cleaned to provide high purity and and Knowles 1985b). Additional introductions were germination levels. Seed viability is retained for a received in the 1920s, but the value of the plant was number of years when seeds are stored under ware- not recognized until the 1930s (Weintraub 1953). house conditions. Intermediate wheatgrass is adapted to areas in the Seeds should be planted at depths of 0.5 to 1.5 inches Northwest, the Intermountain West, and the Plains (1.3 to 3.8 cm) using conventional drills. They germi- States. Both do well as understory species on well- nate quickly when seedbed conditions are favorable drained dry sites in open stands of ponderosa pine and and produce vigorous seedlings. Small seedlings and antelope bitterbrush. At low and midelevations, inter- young plants are able to compete favorably with most mediate wheatgrass is adapted to open parks and other species. Unstable soils and weedy sites are often extremely hot, well-drained sites on exposed slopes. drill seeded with intermediate wheatgrass to provide Intermediate types are more shade tolerant than the permanent stands and uniform, competitive cover in 3 pubescent types and they are better adapted to cooler to 5 years. Seeding at rates below 2 pounds per acre and wetter situations. They perform better on upland (2.2 kg per ha) is recommended when planting mix- benches occupied by Wyoming big sagebrush and in tures for range and wildlife habitat improvement. mountain big sagebrush, herblands, mountain brush, Rates from 8 to 12 pounds per acre (9 to 13 kg per ha) aspen, and fir communities receiving annual precipi- are suggested for planting highly erodible roadways tation of 14 inches (350 mm) or more. Pubescent forms and watershed disturbances. Intermediate wheatgrass are more tolerant of sites with long, warm summers is one of the most successful grasses to establish from than are intermediate forms. They are better adapted broadcast seedings on sites inaccessible to drill seed- to blackbrush, southern desert shrub, and some Wyo- ers. Patches or clumps normally develop when sites ming big sagebrush communities receiving less than are broadcast seeded followed by chaining or harrow- 12 inches (300 mm) of annual precipitation. Popula- ing; these can coalesce over time. tions of intermediate wheatgrass seeded at low eleva- Intermediate wheatgrass is often seeded in mix- tions will eventually shift to support individuals with tures with other herbs to provide forage and cover, but pubescent characteristics. The reverse will occur if it is highly competitive and can suppress and replace pubescent forms are planted in upland communities. many other species. Monsen and Anderson (1993) Intermediate wheatgrass is less drought tolerant than inventoried a 50-year old species trial in central Idaho standard crested wheatgrass, but it is equally cold and found that intermediate wheatgrass and sheep tolerant. Pubescent forms withstand high summer fescue were the two species best able to persist and temperatures of the blackbrush and southern desert displace other introduced and native grasses seeded in shrub types better than standard or fairway crested adjacent plots. wheatgrass. Intermediate wheatgrass is adapted to soil types Uses and Management ranging from heavy-textured with low pH to coarse- textured granite materials with neutral or slightly Prior to about 1950, intermediate wheatgrass was acidic pH. Both the intermediate and pubescent forms not planted as frequently as the crested wheatgrasses are suited to droughty infertile soils, but the pubes- and smooth brome in the Great Plains and Western cent forms are better adapted to saline and alkaline regions of the United States. In the Intermountain soils. area, it has gained importance and is now widely planted. It has been included in seed mixtures for big

328 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses sagebrush, pinyon-juniper, mountain brush, ponde- and others 1968; Hyder and Sneva 1963). Austin and rosa pine, aspen, spruce-fir, and open park herblands. others (1994) found that ‘Luna’ pubescent wheatgrass Because it is a sodformer that forms a dense root received greater preference than other commonly mass (Hafenrichter and others 1968), intermediate seeded grasses when grazed in spring and fall by deer. wheatgrass is used to improve ground cover and to Differences in palatability between cultivars of the stabilize erodible soils and watershed disturbances. It two grasses are due in part to differences in their is adapted to infertile soils associated with roadway phenological development. Some intermediate wheat- disturbances, mined lands, waterways, and other un- grass cultivars mature later in the season and are stable sites. It has become one of the most important more selectively used as pubescent growth forms be- soil conservation species currently seeded. Most col- come dormant (Cook and others 1956). In general, lections exhibit good seedling vigor, establish quickly, intermediate wheatgrass cultivars are more palatable and spread rapidly, even during the first year. Plants to livestock for a longer grazing period than are pubes- remain green and provide effective ground cover cent wheatgrass cultivars. throughout the growing season. Hafenrichter and oth- Intermediate wheatgrass has been seeded in oak ers (1968) reported that intermediate wheatgrass pro- brush and mountain brush thickets. It is tolerant of duces better ground cover than smooth brome in spring, some shade, and provides considerable herbage as an fall, and winter because it does not freeze as easily. It understory species. It begins growth earlier than the is a highly persistent species that retains vigor and shrubs, and has been seeded to suppress shrub growth density and performs better than orchardgrass and and provide more open communities. Intermediate timothy when planted in mixtures on infertile soils. wheatgrass withstands burning and can be used to Intermediate wheatgrass plants spread vegetatively, seed woody sites that tend to burn frequently. Once forming a dense, competitive sod. They are capable of established, it is highly persistent and capable of increasing areas of occupation even when grown on withstanding underground foraging by rodents. harsh sites. Cultivars of intermediate wheatgrass Although intermediate wheatgrass provides useful usually produce a more aggressive and rapidly devel- herbaceous cover in aspen, mountain brush, pinyon- oping sod than do pubescent wheatgrass cultivars. juniper, and sagebrush communities, this species can When grown at low elevations and in more arid situa- slowly replace native herbs and shrubs. Monsen and tions, intermediate wheatgrass forms patches that others (1996) found that extensive seedings in aspen may be widely spaced. Although pubescent wheat- and associated open parks in central Utah have re- grass cultivars respond similarly, they are more likely placed important broadleaf herbs, big sagebrush, an- to form a continuous stand on such sites. When grown telope bitterbrush, and other associated species. Com- at mid and upper elevations, intermediate wheatgrass plete conversion may require as much as 20 to 30 develops a very complete and solid ground cover. Few years. This grass should not be seeded in areas where interspaces remain, and the plants are very competi- native species should be retained. Problems created by tive. Under these conditions, intermediate wheat- seeding intermediate wheatgrass, particularly in as- grass spreads rapidly and can dominate and displace pen and open park herblands, are quite serious. Re- pubescent wheatgrass. moval of wheatgrass from such stands is difficult, and Intermediate wheatgrass is commonly seeded in reestablishment of diverse communities is often not irrigated and dryland pastures. It is suitable for seed- practical. This plant can restore ground cover and ing with alfalfa for hay production or rotational graz- compete effectively with weeds, but it should not be ing. Plants are leafy, but not rank. They are generally planted in areas where retention or reestablishment in the early stage of flowering when alfalfa is ready to of native species is desired. It is particularly restric- cut. At this stage, both species provide high-quality tive to the establishment and survival of shrub seed- herbage. Annual clipping does not weaken stands of lings. It is widely grown for pasture and hay crops from this grass, and plants are capable of recovery and Nebraska to Manitoba, Canada, the Pacific North- regrowth from early spring and summer clippings. west, and to a lesser extent the Intermountain West. Pastures and even dryland seedings can be maintained with considerable grazing. Varieties and Ecotypes Intermediate wheatgrass matures 2 to 3 weeks later in summer than crested wheatgrass, and it provides Cultivars of pubescent and intermediate wheat- much better herbage in mid and late summer (Cook grass forms differ in growth and utility; individual and others 1956). It remains green in summer and fall cultivars should be carefully selected. Varieties of both in mountain brush communities and at higher eleva- forms will grow on the same sites and frequently tions. Both pubescent and intermediate wheatgrass hybridize, but there are some major differences be- are palatable, but investigators have reported sea- tween them. The first promising cultivar of intermedi- sonal differences (Cook and others 1956; Hafenrichter ate wheatgrass was developed through selection from

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 329 Chapter 18 Grasses introduction ‘PI 98568’, received from Maikop, U.S.S.R., The South Dakota Agricultural Experiment Station in 1932 (Hanson 1965). The South Dakota Agricul- released Oahe intermediate wheatgrass in 1961. It tural Experiment Station later released this accession was selected from a Russian introduction, PI98568, as ‘Ree’. Additional cultivars developed from this ac- that also provided material for the Greenar and Ree cession include ‘Chief’, ‘Greenar’, ‘Oahe’, and ‘Slate’. releases. Oahe was developed from two cycles of selec- Some early cultivars including ‘Amur’ have been re- tion for high seed production, forage production, and placed by other, better-adapted cultivars. Only the rust resistance. It was developed for hay, pasture, and most used cultivars are described here. conservation plantings in the Northern Great Plains (Alderson and Sharp 1994). Oahe demonstrates some Intermediate Wheatgrass Varieties of the same vegetative traits as Greenar, but it is more drought tolerant. Both cultivars establish quickly and Chief was released by Agriculture Canada in 1961 develop a dense sod. Oahe does not provide as much as a selection from the cultivar Ree. It was developed late summer and fall herbage as Greenar. Both culti- by mass selection of plants with high seed yields and vars are widely adapted for conservation and pasture good seed quality. It is primarily used in grass and plantings in the Intermountain region. alfalfa pastures and for hay production in the Cana- ‘Reliant’ was selected at the Agricultural Research dian Prairie Provinces. Forage and seed yields are Service Northern Great Plains Research Laboratory slightly higher than those of Ree (Alderson and Sharp in Mandan, ND, and cooperatively released by the 1994). This cultivar has not been widely used in the Agricultural Research Service, Natural Resources Intermountain region. Conservation Service, and the North Dakota Agricul- ‘Clarke’ was released in 1980 by Agriculture Canada tural Experiment Station in Fargo, ND. It was seas a 20-clone synthetic. It was developed through eight lected for its vigor and resistance to leaf-spotting cycles of recurrent selection to combine winter hardi- disease. Reliant is adapted to hay and pasture uses in ness, drought tolerance, seedling vigor, resistance to the Northern Great Plains of the United States and aphid-virus infection, seed quality, and forage and Canada. It has not been extensively planted in the seed production. It has been useful for hay and pasture West, but appears useful in dryland pastures seeded production in dryland or irrigated fields in the Cana- with alfalfa. dian Prairie region and the Northern Great Plains. It ‘Rush’ intermediate wheatgrass was released in is extremely drought tolerant and winter hardy, and 1994 by the Natural Resources Conservation Service produces excellent seed yields (Alderson and Sharp Plant Materials Center, in Aberdeen, ID. It was origi- 1994). Clarke is primarily used in irrigated pastures nally received as A. junceum. Rush develops an exten- in the Western United States. sive sod that provides useful ground cover. It is recom- Greenar is a leafy and highly productive variety that mended for conservation plantings to control erosion was developed at the Natural Resources Conservation on roadways, mine disturbances, and dry waterways Service Plant Materials Center in Pullman, WA, from (Alderson and Sharp 1994). Rush establishes well PI 98568, a collection of the Westover-Enlow expedi- from direct seedings and spreads quickly. It is quite tion to the U.S.S.R. It was released in 1945 as P-2327 competitive and can restrict the occurrence of other and renamed Greenar in 1956 (Alderson and Sharp species. It has an upright growth habit and provides 1994). This release is tall, leafy, productive, medium to considerable spring, summer, and fall forage. late maturing, and mildly sodforming (Hafenrichter The Nebraska Agricultural Experiment Station and and others 1968). Plants are broadleaved and green to Agricultural Research Service cooperatively released dark green. Greenar is widely seeded on rangelands, Slate intermediate wheatgrass in 1969. It was devel- pasturelands, and hayfields in the West. It is used in oped by blending equal quantities of seed from two conservation plantings because it produces consider- unrelated strains, ‘Nebraska 50’ and a derivative of able ground cover and develops an extensive and Amur (Alderson and Sharp 1994). persistent sod. Greenar establishes well and is adapted Slate is an upright variety with large, broad, light- to harsh, dry, exposed sites. It is not shade tolerant. green or blue-green leaves that produces a strong, Plants are disease resistant and begin growth in early spreading sod. It is primarily seeded in mixtures with spring. They retain green herbage late into the season. other grasses and alfalfa in irrigated or nonirrigated Greenar is well adapted to big sagebrush communities pastures and hayfields in the Central Plains. Al- as well as higher elevation sites in aspen and moun- though it has not been seeded extensively in the West, tain herblands. It has persisted well on such sites, it is a useful cool-season forage or hay crop. It is not even when grazed heavily by livestock. It remains tolerant of saline or alkaline soils. green and palatable through the summer and fall The Idaho and Washington Agricultural Experi- months. ment Stations in Aberdeen, ID, and Pullman, WA, and

330 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses the Natural Resources Conservation Service Plant Luna is a tall, highly productive cultivar that is Materials Centers in Aberdeen, ID, and Pullman, WA, widely seeded to furnish spring, summer, and fall released ‘Tegmar’ intermediate wheatgrass in 1968. It herbage. Basal leaves are moderately pubescent, but was developed from accession ‘PI 109219’ collected the seedheads are usually glabrous. Its ability to near Bolu, Turkey, in 1934 (Alderson and Sharp 1994). remain green throughout the summer contributes to Plants are erect and rather small statured. They are its popularity. It is adapted to open parks, aspen, about one-third to one-half the height of the most spruce-fir, mountain brush, and juniper-pinyon wood- robust forms of intermediate wheatgrass. Leaves are lands. It is not shade tolerant, but it grows well on light green to blue green with some hairs along the open, exposed slopes, particularly in areas with warm margins. Tegmar is one of the most commonly used summer temperatures. Because of its drought toler- cultivars in the Western States for stabilizing dry ance, Luna is adapted to big sagebrush and southern waterways, highly disturbed watersheds, and dis- desert shrublands. It requires 10 to 12 inches (250 to turbed rangeland sites. Plants establish quickly and 300 mm) of annual precipitation, but it has been found develop a dense, vigorous, rapidly spreading, and to persist in areas where rainfall is considerably lower. persistent sod that can reduce erosion and withstand Luna seedlings establish quickly and are quite vig- some burial. Tegmar is adapted to warm and low- orous. Plantings reach maturity in 1 to 2 years, and elevation sites. It has been seeded in lower mountain provide some control of spread by annual weeds. Luna brush, big sagebrush, and salt desert communities. is commonly seeded as a dual-purpose species to con- Plants furnish useful spring forage, but other culti- trol erosion and provide forage. Luna is planted on vars may produce more succulent herbage in mid and burned sites, roadways, and unstable watersheds to late summer. Tegmar is frequently seeded alone on provide initial ground cover and supply forage or disturbed sites, particularly on areas with diverse habitat. It provides a productive cover during periods topographical conditions and infertile soils. of drought and heavy grazing. Luna is competitive and Tegmar is not commonly seeded as a hay or pasture can exclude native species. plant on irrigated sites because it does not provide late Topar pubescent wheatgrass was developed from ‘PI summer herbage. It responds poorly to clipping, and it ID7330’ and cooperatively released in 1953 by the is not as productive as other cultivars. It grows well in Washington, Idaho, Oregon, and California Agricul- open stands with big sagebrush and other shrubs, but tural Experiment Stations, and by the Natural Re- it is not shade tolerant. Once stands are established, it sources Conservation Service Plant Materials Centers persists well, even when heavily grazed. at Aberdeen, ID; Pleasanton, CA; and Pullman, WA. Topar is typical of all pubescent and most intermedi- Pubescent Wheatgrass Varieties ate cultivars in being a late-maturing, vigorous, sodforming selection, adapted to droughty sites ‘Greenleaf’ was developed at the Agriculture Canada (Alderson and Sharp 1994). It is seeded in aspen, Research Station in Lethbridge, Alberta, by screening mountain brush, pinyon-juniper, big sagebrush, and open-pollinated progenies for seedling vigor and ten- salt desert shrublands. Seedlings establish quickly dency to creep. Greenleaf is a winter-hardy forage and are very competitive. These characteristics con- cultivar intended for pasture and hay production. It tribute to use of this cultivar on disturbed, weedy, produces bright-green pubescent foliage, and it is infertile sites. Plantings can be used to control weeds fairly tolerant of saline soils and droughty sites. and furnish late-season herbage. Topar is also planted Greenleaf is used primarily for short-duration pasture to control erosion and stabilize watersheds, roadways, forage in the Canadian Prairie region and the North- and waterways. It has been seeded in mixtures to reveg- ern Great Plains. It is similar to ‘Mandan 759,’ but it etate rangeland disturbances. Topar requires between produces more vigorous seedlings. Greenleaf is supe- 10 and 12 inches (250 to 300 mm) of annual rainfall. It is rior to ‘Topar’ in forage production, seedling vigor, and sufficiently drought tolerant for seeding spring and fall winter hardiness (Alderson and Sharp 1994). foothill ranges in the Intermountain area. The New Mexico Agricultural Experiment Station Topar is commonly seeded with Tegmar intermedi- and Natural Resources Conservation Service Plant ate wheatgrass on well-drained, droughty exposures, Materials Center at Los Lunas, NM, released Luna where native vegetation has been lost as a result of pubescent wheatgrass in 1963. It was developed from grazing, wildfires, or other disturbances. Both grasses materials collected by the Westover-Enlow expedition are capable of establishing on soils that dry quickly to the former U.S.S.R. and Turkey in 1934. Luna’s and may be subjected to prolonged periods of summer primary use has been for irrigated pastures and reveg- drought and high temperatures. Both have been used etation of range and watershed disturbances (Alderson to colonize and stabilize unstable soils. Young seed- and Sharp 1994). It has become one of the most lings are able to establish on erosive surfaces, where commonly seeded cultivars in the West. they stabilize the soil and suppress invasive species.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 331 Chapter 18 Grasses

Agropyron sibiricum Most accessions were apparently grouped within the broad crested wheatgrass category. Siberian Crested Wheatgrass _____ Siberian crested wheatgrass is more tolerant of Synonyms drought and alkali soils than is standard crested wheatgrass. Consequently, it performs better at lower Agropyron fragile elevations in big sagebrush and salt desert shrublands. In the Intermountain area it is adapted to upper Description benchlands supporting mountain brush and pinyon- juniper communities and to the extensive Wyoming Siberian wheatgrass is an introduced perennial big sagebrush and shadscale communities bunchgrass that is similar in appearance to both Siberian crested wheatgrass is ecologically better standard and fairway crested wheatgrass, but it has a adapted to arid shrublands of the Intermountain area much narrower spike (fig. 1). It is usually smaller in than standard or fairway crested wheatgrass. It grows stature and develops narrower bunches with less foli- in soils ranging from well-drained and sandy to heavy- age. Its leaves are finer and more decumbent. The textured clay loams. Young plants are able to establish spike is narrow, cylindrical, and usually about 2.3 to on soils that dry quickly in spring and remain dry 4.7 inches (6 to 12 cm) long. The spikelets are some- throughout the summer. This plant demonstrates what spreading and develop directly from the rachis. tolerance to consecutive years of drought when planted Glumes and lemmas are awnless or with awns 0.04 to in the lower regions of the big sagebrush zone. It is 0.08 inch (1 to 2 mm) long (Hitchcock 1950). capable of surviving the hot, dry summers of warm Siberian wheatgrass is part of the crested wheat- desert communities. It is not shade tolerant and gen- grass complex that consists of a series of diploids, erally grows better in regions receiving less than 12 to tetraploids, and hexaploids (Dewey 1974). Diploids 14 inches (300 to 350 mm) of annual precipitation. are represented primarily by the Fairway cultivar. The most commonly used tetraploids are standard Plant Culture crested wheatgrass and Siberian crested wheatgrass cultivars. Selection and hybridization have been ac- Annual seed production of Siberian crested wheat- complished within ploidy levels, but are primarily grass is lower than that of fairway or standard crested confined to the diploid and tetraploid populations, as wheatgrass. However, good seed crops can be pro- hexaploid populations are very rare (Asay 1986). duced on both irrigated and nonirrigated fields. Yields Taxonomic changes have confused the identification averaging 200 pounds per acre (225 kg per ha) for of Siberian crested wheatgrass (Dewey 1986). Jones dryland plantings and 400 pounds per acre (450 kg per (1960) placed all crested wheatgrasses with narrow or ha) for irrigated sites are common. Seed can be har- cylindrical spikes in A. sibiricum. Dewey (1986) rec- vested with combines and cleaned with air-screen ommended acceptance of Tsvelev’s (1976) treatment separators. Cleaned seed lots normally are of high of the wheatgrasses, which identifies three species of viability and purity. Seeds are slightly smaller than crested wheatgrass—A cristatum, A. desertorum, and those of other wheatgrasses. A. fragile. Tsvelev’s decision to combine A. sibericium Cleaned seed lots are usually free of large debris, with A. fragile resulted in the acceptance of A. fragile, and the seeds can be metered through most seeding as it is the older of the two names. Some authors equipment. Seeds should be planted between 0.5 to (Arnow 1987) have recently grouped all crested wheat- 0.75 inch (1.3 to 1.9 cm) deep to support germination grasses into one species, A. cristatum. and establishment. Young and Evans (1986a) found that germination characteristics of Siberian wheat- Ecological Relationships and Distribution grass were similar to those of fairway and standard crested wheatgrass. Hyder and Sneva (1963) reported Siberian crested wheatgrass occurs in eastern Rus- that morphological and phenological development of sia, the Caucasus, Western Siberia, and Central Asia Siberian and crested wheatgrass were very similar. (Weintraub 1953). In Central Asia it is distributed However, when grown under rangeland conditions, from the Caspian Sea to Lake Balkhash (Dewey 1986). seed maturation of Siberian crested wheatgrass is In the United States, the first introduction was re- often 7 to 10 days later than for standard crested ceived in about 1906, but the plant was not success- wheatgrass. fully grown until the 1920s (Weintraub 1953). Dewey Seedlings of Siberian crested wheatgrass cultivars (1986) reported that about 50 accessions entered the are less vigorous than those of most fairway or stan- United States between 1906 and 1940. This exceeds dard crested wheatgrass cultivars, but established the number of A. desertorum introductions during this plants are persistent. Seedling establishment strate- period, but published references to it are infrequent. gies differ among the crested wheatgrasses. Siberian

332 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses crested wheatgrass seedlings do not grow as quickly or Varieties and Ecotypes become as large during the first growing season, as the plants adjust to drought by becoming dormant earlier The Idaho Agricultural Experiment Station, Mos- in the season. Young plants may remain small for 1 to cow, and the Natural Resources Conservation Service 2 years if growing conditions are unfavorable. Plants, Plant Materials Centers at Aberdeen, ID, and Pull- however, generally develop satisfactory stands, even man, WA, cooperatively released the cultivar ‘P-27’ in during periods of drought and in areas where few other 1953 (fig. 7). It was developed from material desig- revegetation species are adapted. Plummer and oth- nated ‘PI 108434’, obtained in 1934 from Kazakhstan, ers (1968) ranked seed germination, initial establish- U.S.S.R., and from the Institute of Plant Industry, ment, growth rate, and persistence of this species as Leningrad, U.S.S.R., by the Westover-Enlow expedi- excellent. tion (Hanson 1959). It was evaluated in field and Siberian wheatgrass can be established with limited nursery plantings beginning in 1935. Individual clones seedbed preparation. Seeds must be covered to assure were selected in 1949 and served as a basis for further acceptable establishment; stands can be planted by increase (Alderson and Sharp 1994). P-27 is primarily drilling or broadcast seeding followed by chaining or used in range seedings. Plants are leafy and fine harrowing. Seeding directly into weedy sites should be stemmed. They are characterized by good seedling avoided, but established plants do compete favorably vigor, high seed yields, and drought tolerance. This with annual weeds. Seedlings are quite hardy, and few cultivar has been the primary accession available for succumb to frost. large seedings.

Uses and Management Siberian crested wheatgrass is less widely planted than other crested wheatgrasses, primarily due to its lower productivity. Most rangeland and pasture plantings are conducted to supply forage for grazing animals, and other, more productive, species are usually planted. Siberian wheatgrass is more drought resistant than most introduced grasses, and it is better able to establish and provide a ground cover in drier habitats. It provides useful, more consistent spring and fall herbage on drier sites than other crested wheatgrasses. It is also a formidable competi- tor with annual weeds. Siberian wheatgrass can be seeded at low rates with bottlebrush squirreltail and Sandberg bluegrass to stabilize Wyoming big sagebrush and salt desert communities. Seedlings develop slowly, allowing establishment of big sagebrush and other shrub seedlings. Siberian wheatgrass is adapted to soil disturbances; it is seeded on roadway and mine disturbances in arid regions. It is often seeded along dry streambeds where soil stabilization is required. Leaves of Siberian crested wheatgrass are usually finer and remain green later into the summer than those of other crested wheatgrasses. Siberian crested wheatgrass is seeded with Russian wildrye to provide later spring and summer pastures. It is also seeded on disturbances in pinyon-juniper and big sagebrush communities to restore forage for big game and domes- Figure 7—‘P-27’, a leafy and fine- stemmed cultivar of Siberian wheatgrass, tic livestock. Although it is somewhat less competitive was released in 1953 and has been widely than either standard or fairway crested wheatgrass, it used (photo courtesy of Loren St. John, is not compatible with native herbs and shrubs, and USDA Natural Resources Conservation should not be seeded in areas where recovery of native Service, Aberdeen Plant Materials Cen- communities is desired. ter, Aberdeen, ID).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 333 Chapter 18 Grasses

‘Vavilov’ Siberian wheatgrass was released in 1994 while the underside is usually smooth. Green leaves by the Agricultural Research Service, the Utah Agri- are flat to involute; when dry they remain stiff and cultural Experiment Station, and the Natural Re- erect, but the edges curl toward the midrib. Auricles sources Conservation Service in Aberdeen, ID. It was are often well developed. Ligules are about 0.04 inch (1 named to acknowledge the contribution of the N. I. mm) long. Spikes are stiff, erect, and about 2.8 to 6 Vavilov Research Institute of Plant Industry, St. Pe- inches (7 to 15 cm) long with one or two spikelets per tersburg, Russia, to the range grass plant breeding node. Spikelets are to 0.4 to 0.8 inch (1 to 2 cm) long and program of the Agricultural Research Service. Vavilov overlap each other by about one-half their length. was derived from accessions obtained from the Vavilov Glumes are firm, unequal, linear-lanceolate to lan- Research Institute; the Stavropol Botanical Garden, ceolate, and gradually tapering from the base to a Stavropoal, Russia; the Eskisehir Plant Breeding Sta- short awn. The first glume is 0.25 to 0.45 inch (6 to 12 tion, Eskisehir, Turkey; and selections from the culti- mm) long and the second 0.3 to 0.6 inch (7 to 15 mm) var P-27 (Asay and others 1995b). The parental acces- long. Lemmas are 0.04 inch (1 mm) long and firm with sions were selected for their retention of green color obscure nerves and a pointed tip or short awn (Arnow and vegetative vigor in late summer. Breeding popu- 1987; Asay 1995a,b; Hitchcock 1950; Sarvis 1941; lations were also screened for seedling vigor, seed Stubbendieck and others 1992). production, plant type, and response to drought, disease, and insects. Vavilov spikes intergrade from the Ecological Relationships and Distribution long narrow forms typical of Siberian crested wheatgrass to shorter, wider spikes more typical of standard Western wheatgrass is morphologically similar to crested wheatgrass (Asay 1995a,b). A tetraploid, thickspike wheatgrass, but it differs in having asym- Vavilov is fully interfertile with standard crested metrical glumes (Cronquist and others 1977). West- wheatgrass and the Hycrest cultivar. ern wheatgrass has recently been placed in a separate Vavilov is a particularly important cultivar because genus and renamed Pascopyrum smithii (Rydb.) Love of its seedling vigor. It is comparable to Hycrest and (Dewey 1984; Love 1980). P. smithii is the sole member consistently better than P-27 in terms of seedling of this genus. Dewey (1975) considers its putative emergence and vigor. Vavilov is also more productive parents to be thickspike wheatgrass and beardless than P-27 (Asay and others 1995b). It is recommended for seeding semiarid sites at elevations below 6,800 ft (2,100 m) that receive 8 to 17 inches (200 to 450 mm) of annual precipitation.

Agropyron smithii Western Wheatgrass or Bluestem Wheatgrass______Synonyms Elytrigia smithii Elymus smithii Pascopyrum smithii

Description Western wheatgrass is a native, rhizomatous, long- lived, cool-season perennial that produces an open and highly uniform sod (fig. 8). Stems are erect and 12 to 24 inches (30 to 60 cm) tall. Leaves are typically blue green, but the entire plant may have a grayish-blue cast that varies among populations. Leaf sheaths are normally glaucous and rough. Leaf blades are about 4 to 10 inches (10 to 25 cm) long and mostly 0.04 to 0.08 inch (2 to 4 mm) wide. They are usually firm, stiff, and Figure 8—A comparison of growth forms of west- taper to a sharp point. The upper surface of the leaf is ern wheatgrass demonstrates some of the varia- rough and distinctly ridged or grooved along the nerves tion in leafiness in this species (RMRS photo).

334 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses wheatgrass. Its adaptability to heavy saline and alka- and 300 pounds per acre (168 to 336 kg per ha) are line soils and its forage qualities reflect the input of expected from cultivated fields with irrigation (USDA these parental species (Dewey 1984). Natural Resources Conservation Service 2002). Western wheatgrass occurs across the western two- Yields from dryland fields are half those of irrigated thirds of Canada and the United States, from Alaska fields. Fields require regular tillage to maintain and British Columbia to Quebec, and southward proper spacing and prevent overcrowding by sod- through the Western and Central United States (Arnow ding. Plants flower in June, and seeds ripen in August 1987; Asay 1996) east of the Cascade Mountains to or September (Wambolt 1981), later than for many New Mexico (Hanson 1972), the Central and Northern other grasses. Seed stalks are erect and project above Great Plains, and the Texas Panhandle (Hanson 1972; the leaves, facilitating combine harvesting. Seeds are Hitchcock and others 1969). It is the only octoploid large, firm, and easily separated from the spikelet wheatgrass native to North America (Asay 1995a,b). using standard cleaning equipment. Ecotypes of western wheatgrass varying in stature, Western wheatgrass seeds, particularly those from color, and sodding traits occur in a variety of plant wildland collections, do not germinate readily or uni- communities. On the Central Great Plains, it is abun- formly, resulting in erratic establishment (Beetle 1955; dant in lowlands, and may occur as nearly pure stands Ferguson and Frischknecht 1985; Monsen and aligning watercourses (Sarvis 1941). Throughout this McArthur 1985). The seeds often exhibit low or de- region it grows with blue grama, sideoats grama, alkali layed germination at warm temperatures between 66 sacaton, buffalo grass, and needlegrasses. In the West- to 73 ∞F (19 to 23 ∞C) (Brown and Hallman 1984; ern United States it is often the dominant grass in salt Plummer 1976), but alternating temperatures of 59 to desert, big sagebrush, mountain brush, and pinyon- 80 ∞F (15 to 30 ∞C) tend to promote germination. juniper communities. In Utah it rarely occurs on slopes Erratic germination decreases survival in rapidly above 6,900 ft (2,130 m) (Arnow 1987), but it grows at drying seedbeds. Although slow- or late-developing higher elevations in Wyoming and Colorado (Dittberner grass seedlings may succumb to competition with and Olsen 1983). Western wheatgrass often grows in weeds, western wheatgrass seedlings are extremely association with bluebunch wheatgrass and needle- drought tolerant. When seeded in mixtures with more and-thread on upland benches and with bottlebrush aggressive species, poor and spotty stands may de- squirreltail and Sandberg bluegrass in Wyoming big velop. Increasing the seeding rate usually does not sagebrush and salt desert communities. improve plant density. However, when seeded in weed- Beetle (1955) ranked this species as the most alkali free, but harsh environments typical of mine distur- tolerant of all North American wheatgrasses. It is bances, this grass establishes satisfactorily. Seed ger- commonly found on heavy clay soils with moderate mination and establishment characteristics of recently alkalinity in salt desert shrublands. It is common in developed cultivars are much improved over previ- cold desert uplands of northern Utah, western Wyo- ously used wildland collections. ming, and eastern Idaho where salty outcrops occur Established plants are vigorous and drought resis- with Wyoming big sagebrush, winterfat, and various tant. Plants spread vegetatively from rhizomes, form- saltbush species. It is also common in dry valley ing open, irregular patches, even when seeded alone. bottoms, swales, and dry drainages where periodic Western wheatgrass can be seeded with slower devel- flooding occurs (Rogler 1973). Although western wheat- oping native species, including shrubs. grass exists on fine-textured soils, it grows on well- drained bottomlands, open plains, and benchlands of Uses and Management lower mountain and foothill ranges in the Intermoun- tain region (Stewart and others 1939). It occurs in Western wheatgrass is one of the most important areas receiving 10 to 20 inches (250 to 500 mm) of native forage grasses of the Northern Great Plains annual precipitation, but performs well in the 10- to (Sarvis 1941) and the Intermountain West (Stewart 14-inch (250- to 360-mm) zone. It is sometimes an and others 1939; Vallentine 1961). It has not been important understory species in ponderosa pine and planted as extensively on range and wildland sites as pinyon-juniper, but it is not shade tolerant and is crested wheatgrass and intermediate wheatgrass be- usually restricted to openings between trees. cause its seedling establishment is less reliable and its herbage yields are lower. Western wheatgrass begins Plant Culture growth relatively early in spring, and plants mature late in the season. Standing plants cure well, remain- Seed production of native western wheatgrass stands ing palatable and nutritious. Energy values are rated in the Northern Great Plains is generally greater than good (Dittberner and Olson 1983). Livestock and game from stands in more arid regions. Released cultivars graze it during most seasons. Because it provides good have improved seed production capabilities compared winter grazing, it is often managed to provide late fall to wildland collections. Yields ranging between 150 and winter pastures (Sarvis 1941).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 335 Chapter 18 Grasses

Western wheatgrass has been reduced or eliminated cannot be harvested from wildland stands to plant from many Western rangelands by continuous and large projects. Consequently, for most seedings, the prolonged grazing. Close and repeated clipping can best adapted of the currently available cultivars must reduce yields and weaken stands (Wasser 1982). How- be selected. ever, plants have demonstrated the ability to recover The Natural Resources Conservation Service in Los if protected from grazing. Plants spread quickly from Lunas, NM; the Colorado Agricultural Experiment rhizomes and are able to repopulate abandoned lands Station in Fort Collins; and the New Mexico State (Rogler 1973; Weaver and Albertson 1956). Consider- Highway Department in Santa Fe, cooperatively re- able recovery has been noted in Wyoming big sage- leased ‘Arriba’ western wheatgrass in 1973. It was brush and pinyon-juniper communities protected from developed from a collection obtained in 1957 from a grazing for 1 to 20 years. Plants have increased during site near Flagler, CO, at an elevation of 4,970 ft (1,530 periods of drought (White and others 1978) and re- m) that receives annual precipitation of nearly 16 placed annual weeds. Sarvis (1941) reported that inches (400 mm). Seed production was increased by western wheatgrass recovers so quickly in selection through five generations (Alderson and Sharp noncultivated fields that it has been called “go-back” 1994). Seeds of Arriba germinate rapidly and uni- grass. Rate of recovery is based on plant vigor, density, formly, providing vigorous seedlings. Seed production and the presence of other species. of this cultivar exceeds that of other tested accessions. Western wheatgrass is particularly important for Plants establish quickly and spread rhizomatously. restoring semiarid communities because it is one of Herbage yields are good, but less than those of the the few species adapted to these sites and it can be variety ‘Barton’ when grown in pastures. Arriba has established successfully. It grows in association with performed well on arid mine sites. It has also done well other native species on sites that have been seriously when seeded on semiarid shrublands. It produces altered and are occupied by annual weeds. It can be dark green foliage that is sought by grazing animals. seeded with bluebunch wheatgrass, slender wheat- Young plantings establish quite well, even on infertile grass, bottlebrush squirreltail, Sandberg bluegrass, sites. Arriba and ‘Rosana’ are the two most widely and related broadleaf herbs to reestablish native com- seeded cultivars of western wheatgrass in the Inter- munities and promote secondary succession. mountain region. Western wheatgrass has been used to revegetate Barton western wheatgrass was cooperatively re- mine disturbances. Although establishment from di- leased in 1970 by the Natural Resources Conservation rect seeding is slow, plants spread by rhizome develop- Service Plant Materials Center at Manhattan, KS; the ment and stabilize exposed soils (Thornburg 1982). Kansas Agricultural Experiment Station, and the This grass is particularly useful for controlling erosion Agricultural Research Service. The original seed was of sandy soils (Scheetz and others 1981) and for provid- collected from native grassland in Barton County, KS. ing protective ground cover throughout the entire It is used in the Midwest to stabilize disturbed soils. It year. It establishes quickly on barren mine wastes is the most productive cultivar currently available that are free of competitive weeds. Established sod is (Alderson and Sharp 1994), and it is also strongly very persistent, but not restrictive to seedling inva- rhizomatous. It has not been extensively seeded in the sion by some native herbs and shrubs. Intermountain area. Western wheatgrass is also commonly planted on The Nebraska Agriculture Experiment Station and saline seeps associated with mine disturbances (Rogler the Agricultural Research Service cooperatively re- 1973). It grows well on loam and clay loam soils and leased ‘Flintlock’ in 1975. It is a broad-based cultivar fine-textured mine wastes, and it persists on soils with genetically, derived from materials collected in cen- moderate alkalinity (Reitz and Morris 1939; Wambolt tral and southwestern Nebraska and northwestern 1981), overflow sites, and subirrigated wetlands with Kansas. Flintlock produces aggressive rhizomes. It is poor drainage. It also withstands short periods of commonly used for conservation plantings, dryland flooding. hay production, and early-season pastures in the Cen- Western wheatgrass recovers from wildfires and tral Great Plains (Alderson and Sharp 1994). prescribed burning through rhizome proliferation ‘Rodan’ was developed from materials collected in (Wasser 1982; White and Currie 1983). Aboveground the Missouri River bottoms near Mandan, ND. The stems may be burned, but little heat is transferred to Agricultural Research Service, the Natural Resources the root meristematic tissue (Gartner and others 1978). Conservation Service, and the North Dakota Agricul- ture Experiment Station released it in 1983. Rodan is Varieties and Ecotypes primarily adapted to the Great Plains of the United States and the prairies of Canada. This cultivar was Considerable variability exists among populations selected for its seed production and drought tolerance. of western wheatgrass, and use of local or adapted It is used primarily for revegetation of disturbed materials is recommended. However, sufficient seed lands. It is also planted for grazing and hay production

336 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses because it yields well on dry sites (Alderson and Sharp mostly 0.4 to 0.8 inch (1 to 2 cm) long. Those near 1994). midlength on the spike rarely exceed the internodes by The Natural Resources Conservation Service Plant more than one-third their length. Glumes are mostly Materials Center at Bridger, MT, and the Montana unequal, somewhat oblanceolate, three to seven nerved, Agricultural Experiment Station released ‘Rosana’ 0.18 to 0.47 inch (4.5 to 12 mm) long, and obtuse to western wheatgrass in 1972. It was selected from acute or rarely awn tipped. Lemmas are 0.28 to 0.51 materials harvested in native meadows near Forsyth, inch (7 to 13 mm) long, acute or terminating in a widely MT, and developed for seeding irrigated hayfields and divergent awn. pastures. It has also been successfully planted on Both diploid (2n = 14) and autotetraploid (4n = 28) rangelands and mine disturbances (Alderson and Sharp forms occur in the West, but tetraploids are primarily 1994). Seed is readily available. Rosana is one of the limited to eastern Washington, northwestern Idaho most widely used cultivars in the Intermountain re- (Hartung 1946), and Canada. Dewey (1966) concluded gion. Its range of adaptability, however, has not been that bluebunch wheatgrass has been involved in the adequately defined. phylogeny of several other grasses. Beardless wheat- Rosana has been used to control weeds and provide grass has been designated a separate species by some ground cover because it produces a closed sod soon authors, but Heller (1900), Beetle (1961), and Cronquist after establishment. Seedlings are vigorous and estab- and others (1977) combined it with bluebunch wheat- lish well, particularly when seeded in mixtures or grass. Beardless wheatgrass lacks the distinctive di- planted on harsh sites. Moderate to good amounts of vergent awn, a recessive trait of bluebunch wheat- light blue-green herbage are produced annually. grass (Asay 1995a,b). Stebbins and Fung (1953) ‘Walsh’ was developed from southern Alberta and presented cytogenetic evidence demonstrating that southwestern Saskatchewan collections and released both varieties are completely interfertile. Both hybrid- in 1982 by Agriculture Canada. It is a dual-purpose ize freely and produce fertile progeny. Daubenmire cultivar, adapted for pastures and hay production. It (1960) concluded that the cespitose growth forms of is also extensively seeded to revegetate disturbed lands in Canada. Walsh is tolerant of saline soils, drought, and periodic flooding (Alderson and Sharp 1994).

Agropyron spicatum Bluebunch Wheatgrass ______Synonyms Agropyron inerme Agropyron spicatum var. inerme Elymus spicatus Elytrigia spicata Pseudoroegneria spicata

Description Bluebunch wheatgrass (fig. 9) is a perennial, cool- season, native bunchgrass (Arnow 1987; Hitchcock 1950; Stubbendieck and others 1992). Plants are rarely rhizomatous. Stems are erect and 24 to 40 inches (60 to 100 cm) tall. Sheaths are glabrous or puberulent. Blades are flat or involute, 0.04 to 0.08 inch (1 to 2 mm) wide, and 2 to 10 inches (5 to 25 cm) long. The upper surface normally has small hairs, but the underside is smooth. Auricles are well developed. Ligules are membranous and about 0.04 inch (1 mm) long. Spikes are typically erect, slender, and 2.4 to 8 inches (6 to 20 cm) Figure 9—Bluebunch wheatgrass is one of the long with a continuous rachis. Spikelets are rarely most important native bunchgrasses (photo cour- more than one per node, five to eight flowered, and tesy of Kevin Jensen, USDA ARS, Logan, UT).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 337 Chapter 18 Grasses bluebunch wheatgrass are adapted to arid grasslands Plummer and others (1968) reported that this grass and semideserts, while rhizomatous forms require exhibits considerable variation. Some beardless and more mesic sites. bearded forms have similar areas of adaptation, but The cultivar ‘Secar’ was originally identified and some exist in differing sites. Plummer and others released in 1980 as a bluebunch wheatgrass. Asay (1968) concluded that sod-forming strains are supe- (1995a,b) later referred to it as “thickspike wheat- rior to bunch types for seeding game ranges with grass” because it is morphologically similar to variable topography in Utah, because they are adapted bluebunch wheatgrass, but genomically aligned with to a wider range of moist to dry habitats. Daubenmire thickspike. It is now considered a separate taxon, (1960) noted that caespitose individuals prosper in Snake River wheatgrass (Agropyron spicatum ssp. arid grasslands in eastern Washington and northern laceolata), originally designated Elymus lanceolatus Idaho, and that rhizomatous types dominate in more ssp. wawawaiensis (Carlson 1986), and more recently mesic grasslands. Dewey (1970) suggested that the Elymus wawawaiensis (Natural Resource Conserva- presence of rhizomes may result from introgression tion Service 2002). Snake River wheatgrass, which with thickspike wheatgrass. A greater chance for includes Secar, is most prevalent along the lower introgression would occur at upper elevations where Snake River drainage of central Idaho (Asay 1996). thickspike wheatgrass is more common. Rhizomatous Since Secar was initially considered a variety of types are not always restricted to more mesic condi- bluebunch wheatgrass and as it is ecologically and tions, and moderately spreading growth forms are morphologically aligned with this species, we have sometimes found growing in drier situations. Some treated it as a bluebunch wheatgrass in this text. rather extensive rhizomatous populations occur in pinyon-juniper, antelope bitterbrush, and big sage- Ecological Relationships and Distribution brush communities. Since this species occurs over such a wide range of sites, and exhibits considerable Bluebunch wheatgrass occurs from Alaska to difference in vegetative characteristics, selection and Saskatchewan and south to California, Arizona, New use of specifically adapted ecotypes is recommended. Mexico, and Texas (Arnow 1987). On the Palouse Prairie it frequently occurs in pure stands, but it is Plant Culture most often codominant with Idaho fescue and Sandberg bluegrass (Miller and others 1986). It is often a domi- Seeds of bluebunch wheatgrass cultivars are com- nant species on grasslands of the Pacific Northwest. In monly produced under cultivation. Wildland stands areas of the Intermountain area receiving 7 to 30 normally produce seed crops most years (Stewart and inches (150 to 760 mm) of annual moisture it grows in others 1939), and plants are capable of producing some association with big sagebrush and antelope bitter- seeds even during periods of drought. brush (Stoddart 1941; Wright and Bailey 1982). Plants grown under cultivation are usually planted Bluebunch wheatgrass generally occurs on sagebrush in rows spaced 24 to 36 inches (60 to 90 cm) apart. flats, low foothills, and benchlands (Pickford 1932). It Satisfactory crops can be grown under irrigation or if is not highly shade tolerant, but it does occur in sufficient moisture is available, under dryland condi- pinyon-juniper, mountain brush, aspen, ponderosa tions. If irrigation is required, application is made pine, and spruce-fir forests. It forms distinctive and prior to pollination or the late boot stage. In fields that important associations with ponderosa pine at mid dry rapidly, irrigation soon after harvest may be and upper elevations in Oregon, Washington, Mon- required to ensure fall regrowth and tillering. Under tana, and Idaho, but it is equally important in more cultivation, bluebunch wheatgrass fields normally arid grass or shrub communities on dry, well-drained produce 300 to 385 pounds of seed per acre (336 to 430 slopes. kg per ha) (Hafenrichter and others 1968). Seeds ripen Bluebunch wheatgrass occurs in some unique plant early, generally by mid-July. Moderate amounts of communities. It exists in restricted areas with low fertilizer are required to maintain productivity. sagebrush, alkali sagebrush, and black sagebrush in Bluebunch wheatgrass develops an extensive root Nevada (Zamora and Tueller 1973); pygmy and stiff system, and properly spaced plantings are normally sagebrush in Idaho, throughout the extensive steppes somewhat competitive with weeds. Seeds usually ripen of Washington (Daubenmire 1970); and in the sage- uniformly, but mature seeds shatter quickly. Fields brush-grass steppes of Colorado (Francis 1983) and can be swathed when seed is in the soft-dough stage southern Idaho (Hironaka and others 1983). and then combined; however, direct combining is most Bluebunch wheatgrass occurs on soils that vary in practical. Bluebunch wheatgrass seed is easily cleaned, texture, depth, and parent materials. It is often found but awns must be removed from some cultivars and on coarse, well-drained mountain slopes (Tisdale and wildland collections. Infertile florets can be removed Hironaka 1981). It is not tolerant of alkaline or saline during cleaning. Seed viability from wildland collec- soils, but is does grow on dry, calcareous soils. tions may be less than 60 percent.

338 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Bluebunch wheatgrass is best sown in fall, but it will Seedings of bluebunch wheatgrass and other adapted establish from early spring plantings if moisture con- native herbs are required to restore native shrub and ditions are favorable. Seeds can easily be planted with herb associations. Antelope bitterbrush, Stansbury most drills or other seeders. Seeds are of moderate size cliffrose, and mountain big sagebrush have been elimi- and should be planted at depths of 0.5 to 1.5 inches (1.3 nated from many game ranges through excessive graz- to 3.8 cm). Seedlings are not as vigorous or competitive ing by wild ungulates. In many cases, cheatgrass and as those of cheatgrass (Harris 1967), but they estab- other annual weeds have invaded and become the lish and persist well when planted in weed-free seed- dominant vegetation. Reestablishment of native shrubs beds. Bluebunch wheatgrass seeds germinate fairly by interseeding and transplanting shrubs in clearings uniformly, seedlings grow rapidly, and mature stands within annual grass stands has been successful, but will develop in 2 to 3 years. Young plants develop without reestablishment of the native understory, the extensive root systems, and after about 2 years they seeded shrubs are unable to reproduce naturally. provide considerable competition. Planting success is dependent, in part, on adaptabil- Dewitt (1969) compared the germination rates of 51 ity of the seeded ecotype. Nearly all seedings using accessions of bluebunch wheatgrass in an attempt to bluebunch wheatgrass are conducted using seed of select sources capable of germinating at cooler tem- three available cultivars: ‘Goldar’, Secar, and peratures and competing more successfully with ‘Whitmar’. Although these are useful and widely used cheatgrass seedlings. Although differences among col- cultivars, each has specific areas of adaptation. Goldar lections were noted, they were not great enough to be and Whitmar are better adapted to uplands and most of practical importance. Improvement of germination moist sites, while Secar is better adapted to the lower rate and emergence success through selection of larger ranges, particularly in the Snake River drainage. seeds from different populations appears possible None are universally adapted throughout the entire (Kitchen and Monsen 1994). As additional ecotypes range of this grass. In addition, none of these cultivars are selected for wildland seedings, emphasis on devel- are well adapted to Great Basin sites. As additional opment of seedling vigor and establishment traits releases or seed sources become available, planting should be emphasized. success is likely to improve. Bluebunch wheatgrass has often been replaced by crested wheatgrass in range and wildlife seeding Uses and Management projects. However, bluebunch wheatgrass develops strong stands, and it can be seeded as a major compo- Bluebunch wheatgrass is one of the most important, nent of native mixtures. Established seedings of productive, and palatable native grasses in big sage- bluebunch wheatgrass are very persistent brush communities. It is a major species in sagebrush- (Hafenrichter and others 1968; McGinnies and others grass, pinyon-juniper, and ponderosa pine communi- 1983; Plummer and others 1955). Native stands elimi- ties. It is valuable because of its widespread distribution nated by grazing have been reestablished by direct and because it provides excellent forage in spring, seeding. Monsen and Anderson (1993) found that summer, and fall (Vallentine 1961). It is also used in bluebunch wheatgrass plantings established in south- winter at certain locations. Bluebunch wheatgrass central Idaho in 1927 persisted through two wildfires begins growth in early spring as soil temperatures and prevented invasion of introduced perennial grasses approach 42 ∞F (17 ∞C) and air temperatures reach 58 during a 64 year period. to 77 ∞F (24 to 32 ∞C) (Anderson 1991). Although Bluebunch wheatgrass stands can be restored with differences in palatability occur among ecotypes protective management if some residual plants re- (Willms and others 1980), this grass is palatable to main. Considerable increase in plant density was most classes of domestic stock and big game, and noted on a number of sites in the Intermountain West supplies excellent herbage even after growth has between 1989 and 1994, during a period of extended ceased. It cures well and in many locations it is used as drought. Although bluebunch wheatgrass seedlings winter forage by wildlife (Peek and others 1979). are not able to compete with annual weeds as success- Reestablishment of bluebunch wheatgrass is desirable fully as crested wheatgrass seedlings (Harris 1967), over a broad range of sites. This species has been reduced they are quite hardy. Bluebunch wheatgrass can be from extensive areas by poorly regulated grazing seeded in areas where it once dominated to reduce (Anderson 1991; Blaisdell and Pechanec 1949; Forsling weed encroachment and provide a permanent cover. and Dayton 1931; Rickard and others 1975). Loss of When seeded in disturbances where residual native bluebunch wheatgrass and related species has resulted species still occur, established bluebunch wheatgrass in rapid invasion by annual weeds, particularly cheatgrass usually allows for natural spread and encroachment of (Christensen 1963; Cottam and Evans 1945; Daubenmire other native species. 1940; Pickford 1932). Christensen (1963) reported that mechanical disturbances and frequent fire also reduced

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 339 Chapter 18 Grasses the occurrence of the species. This change in species bluebunch wheatgrass because it has a much greater composition has diminished forage production and spe- capacity to allocate limited carbon resources to foliage cies diversity, and increased fire frequency. As a result of regrowth following clipping or grazing (Caldwell and overgrazing by domestic animals, cheatgrass and other others 1981; Caldwell and Richards 1986b; Richards annual exotic weeds have displaced bluebunch wheat- and Caldwell 1985). grass communities across extensive areas of the West Many bluebunch wheatgrass communities have been (Christensen 1963; Cottam and Evans 1945; Daubenmire converted to crested wheatgrass and intermediate 1940; Pickford 1932). Christensen (1963) reported that wheatgrass seedings because these species withstand mechanical disturbances and frequent fires have also more intensive late spring and summer grazing. Most reduced the occurrence of this species. wildland sites are not managed as intensive pastures, Reestablishment of bluebunch wheatgrass and as- and costs for site conversion usually do not justify this sociated herbs has been hindered by a lack of adapted effort. Plummer and others (1955) found that bluebunch seeds and by the competitive ability of cheatgrass and wheatgrass, including beardless bluebunch wheat- other weeds. Elimination and control of weeds is grass, is more productive than crested wheatgrass on essential if bluebunch wheatgrass communities are to poor sites. These investigators also found that on sites be restored. Although bluebunch wheatgrass seed- receiving more than 10 inches (250 mm) of annual lings and those of many other perennial grasses are precipitation, yields of bluebunch wheatgrass are com- not as competitive as cheatgrass and other annual parable to those of crested wheatgrass, but fluctuate weeds, reestablishment of bluebunch wheatgrass com- less than those of introduced wheatgrasses. Bluebunch munities by careful management and artificial seed- wheatgrass is now widely used to restore native com- ing are the most practical solutions to weed invasion munities. Plantings in the big sagebrush, pinyon- and spread. juniper, and ponderosa pine types are successfully Bluebunch wheatgrass plants are particularly sen- seeded with this species when planted alone or in sitive to season and intensity of grazing. Most plants mixtures. succumb from repeated moderate to heavy clipping in Bluebunch wheatgrass is an important species for late spring at the height of the growing season wildlife. It provides useful forage, and along with (Daubenmire 1940). Hyder (1972) found that species woody shrubs furnishes cover and protection. Bodurtha such as bluebunch wheatgrass produce culms late in and others (1989) found that mule deer preferred the growing season when the apical meristems are bluebunch wheatgrass communities that approximated elevated as internode elongation proceeds. These mer- the composition of climax communities during both istems are susceptible to removal by grazing, and spring and summer grazing in Oregon. Deer and elk further growth must originate from axillary buds at selectively graze Utah pinyon-juniper sites that are the base of the plant. Defoliation during spring, when seeded with bluebunch wheatgrass. vegetative growth is proceeding rapidly and the apical Bluebunch wheatgrass furnishes ground cover and meristem is elevated, is most detrimental (Stoddart soil protection for steep slopes in watersheds where 1946b). Miller and others (1986) concluded that plants erosion is a serious problem. It is particularly impor- can withstand use early in the growing season if tant in semiarid environments where loss of ground conditions permit adequate growth after grazing. cover creates major watershed problems and distur- Rickards and others (1975) found that 50 percent bances are difficult to restore. It provides excellent use for 2 consecutive years on previously protected cover in ponderosa pine and pinyon-juniper wood- sites resulted in a 26 percent decrease in the basal area lands where seasonal storms can generate consider- of bluebunch wheatgrass. Heady (1950) concluded able runoff. that clipping once to a 6-inch (15-cm) height at flower- Bluebunch wheatgrass is fire tolerant and regener- ing was too severe to permit plants to maintain them- ates vegetatively following burning. Only small selves. Wilson and others (1966) found bluebunch amounts of litter accumulate near the base, limiting wheatgrass may be eliminated by grazing to a 1-inch fuel buildup and potential fire damage to root crowns (2.54-cm) stubble height for 3 consecutive years dur- (Antos and others 1983). Leaves and stems are coarse ing the boot stage. Mueggler (1975) found that and burn quickly, transferring little heat to the soil bluebunch wheatgrass plants growing with competi- (Wright 1985). Although bluebunch wheatgrass usu- tion may require 8 years to approach normal vigor ally survives fires, postburn production, ground cover, following late spring clipping. tillering, and seed production vary among burns. Crown Bluebunch wheatgrass plants do not curtail root cover can be reduced over 52 percent following growth following defoliation (Richards 1984). This wildfires (Conrad and Poulton 1966), and foliage reduces the availability of carbon reserves for shoot production normally decreases for 2 to 3 years development (Caldwell 1984; Richards 1984). Crested (Daubenmire 1975b; Muggler and Blaisdell 1958). wheatgrass has often been planted as a substitute for

340 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Varieties and Ecotypes bluebunch wheatgrass (2n = 4x = 28). The Agricultural Research Service, Forage and Range Research Labo- Only three cultivars of bluebunch wheatgrass are ratory, Logan, UT, the Utah Agricultural Experiment currently available. Additional populations or ecotypes Station, and the Natural Resource Conservation Ser- are needed. Individual ecotypes differ in adaptability, vice released this cultivar in 1989. NewHy was derived and local collections should be seeded where available from two germplasms (‘RS-1’) and (‘RS-2’) released in releases are not adapted. 1980 (Asay and Dewey 1981). The initial RS hybrid Anatone bluebunch wheatgrass is a selected population was established in 1962. These plants were germplasm originating from Asotin County, WA, at an only partially fertile, but fertility and growth charac- elevation of 3,200 to 3,600 ft (975 to 1,097 m). It teristics were improved through an eight-generation exhibits good seedling vigor and is recommended for selection process. The primary objectives in the selec- use on bluebunch wheatgrass sites receiving at least tion process were to combine the vigor, productivity, 10 inches (25 cm) of annual precipitation (Monsen and salinity tolerance, and persistence of quackgrass with others 2003). the drought tolerance, caespitose growth habit, and Goldar bluebunch wheatgrass was initially selected seed and forage quality of bluebunch wheatgrass (Asay by the Natural Resource Conservation Service Plant and others 1991). Caespitose types were emphasized, Materials Center at Pullman, WA, but it was moved to but annual rhizome development of NewHy ranges the Plant Materials Center in Aberdeen, ID, where it between 1.6 and 3.3 ft (0.5 and 1.0 m) per year. was released in 1989. Seed was originally collected NewHy can grow in saline soils (Currie and others from an open ponderosa pine community on the 1986); its salt tolerance approaches that of tall wheat- Umatilla National Forest near Asotin, WA, at an grass. NewHy is most productive on slightly saline or elevation of 1,000 to 1,550 ft (310 to 475 m) in 1934. alkaline sites receiving about 17 inches (330 mm) of Goldar was developed by mass selection from spaced annual precipitation. Seed quality varies among lots. plantings (Alderson and Sharp 1994). When seeded alone, a seeding rate 8 to 17 pounds per Goldar is a diploid form of bluebunch wheatgrass, an acre (9 to 19 kg per ha) is sufficient. Poor seed germi- ecotype of A. s. ssp. spicata (USDA Soil Conservation nation can result in weak stands, but as plants de- Service 1989). This is a vigorous cultivar, representa- velop, tillering and rhizome development in the third tive of material found in northeastern Oregon and and fourth year provide more complete cover. NewHy southern Washington. It is a leafy, productive, and provides high-quality forage, comparable to that of densely tufted bunchgrass that produces some tillers intermediate wheatgrass. Plants begin growth in early and short rhizomes. It is relatively drought tolerant spring and remain green until late in the growing when compared with Whitmar. Goldar produces seeds season. On dryland ranges NewHy provides late spring with the typical divergent awns of the species; these grazing and competes well with weeds. It tolerates must be removed prior to seeding. moderate grazing and recovers quickly following use, Goldar is recommended for seeding rangeland sites an improvement over bluebunch wheatgrass. to improve native diversity and to provide wildlife P-7 bluebunch wheatgrass is a multiple-origin habitat and forage for grazing animals. It is adapted to polycross generating by mating 23 open-pollinated, sites receiving 12 to 14 inches (300 to 360 mm) of native-site collections and two cultivars from Wash- annual precipitation. In the Intermountain West, it ington, Oregon, Nevada, Utah, Idaho, Montana, and has done well on pinyon-juniper and upper benchlands British Columbia. P-7 is intended to provide increased occupied by mountain big sagebrush. It has survived genetic diversity upon which natural selection may when seeded in disturbed Wyoming big sagebrush operate within a single germplasm. It is intended for communities receiving at least 12 inches (300 mm) of use on semiarid to mesic sites in the Intermountain annual precipitation. It is not as drought tolerant as Region. P-7 has very good seed yields. Genetically, Secar, but it is much more palatable and remains P-7 corresponds to the “P” metapopulation, which green later in the season. It provides considerable fall includes native populations in southeastern Washing- greenup, and develops a very dense and competitive ton, northeastern Oregon, and western Idaho, the ground cover. Plants cure well and are grazed as region from which the great majority of the P-7 winter pastures. Overseeding Goldar on depleted big components originated (Jones 2002a). sagebrush communities by aerial seeding and chain- Secar was selected at the Plant Materials Center at ing has been very successful. This cultivar provides Pullman, WA, from a collection obtained by J. L. useful herbage in mixed and single-species plantings. Schwendimann near Lewiston, ID, in 1938. Originally It is particularly useful for grazing in spring and early considered a bluebunch wheatgrass, it was later iden- summer. Goldar is sensitive to grazing during the tified as Snake River wheatgrass. The Agricultural vegetative stages of growth. Research Service; the Idaho, Oregon, Montana, and ‘NewHy’ RS hybrid wheatgrass was developed by Wyoming Agriculture Experiment Stations; and the artificially crossing quackgrass (2n = 6x = 42) and

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 341 Chapter 18 Grasses

Natural Resources Conservation Service Plant Mate- Whitmar beardless bluebunch wheatgrass was origi- rials Center in Pullman, WS, released Secar in 1980. nally released as beardless wheatgrass and is awnless Secar is a densely tufted bunchgrass with many fine or nearly awnless. It was released in 1946 by the leaves and stems. Seeds are small and mature early in Washington, Idaho, and Oregon Agricultural Experi- the season. This cultivar is more drought tolerant and ment Stations, and by the Soil Conservation Service better adapted to harsh sites than other bluebunch Plant Materials Centers at Aberdeen, ID, and Pull- wheatgrass cultivars. It is slower to establish and less man, WA. Whitmar was developed from material productive than the more robust cultivars. collected from native Palouse Prairie grasslands near Secar is adapted to low-elevation sites in the Pacific Colton, WA. Original collections were growing at an Northwest receiving 8 to 12 inches (200 to 300 mm) of elevation of 2,780 ft (845 m) on a silt loam soil in an annual precipitation (Alderson and Sharp 1994). Secar area receiving 20 inches (500 mm) of annual precipita- is also seeded in Wyoming big sagebrush communities tion (Alderson and Sharp 1994). on the Snake River Plain of southern Idaho, and in Whitmar is considered an intermediate growth form. northern Nevada and central Utah. It is less produc- It is erect or semierect with moderately abundant, tive than Goldar when seeded on more upland sites, medium-coarse stems and leaves. The cultivar pro- but it produces considerable herbage under arid con- vides good spring and fall herbage and it is a good seed ditions. Secar is well adapted to sites with long, hot producer (Alderson and Sharp 1994). Whitmar is pri- summers, as it enters dormancy in late spring or early marily used in the Pacific Northwest. Although ini- summer. It forms closed stands when seeded in rows tially seeded in the Intermountain area, it has been spaced less than 28 to 30 inches (71 to 76 cm) apart. replaced by Goldar and Secar. Established stands, particularly in low rainfall areas, are somewhat competitive with annual weeds. Secar is fire tolerant and recovers well following wildfires. Agropyron trachycaulum Herbage is not as selectively grazed as other cultivars Slender Wheatgrass ______or ecotypes. Secar establishes well when seeded in mixtures with other natives. Synonyms The ‘SL-1’ germplasm was developed and released Agropyron caninum by the Agricultural Research Service Forage and Range Agropyron latiglume Research Laboratory in Logan, UT, in 1990. It is a Agropyron pauciflorum hybrid derived from the diploid form of bluebunch Agropyron subsecundum wheatgrass and thickspike wheatgrass. The initial F1 Elymus pauciflorus hybrid was made in 1965 by crossing a bluebunch Elymus subsecundus wheatgrass collection obtained in Utah with a Nevada Elymus trachycaulus collection of thickspike wheatgrass (Dewey 1965). The F1 hybrid was completely sterile, but use of colchicine Description permitted development of a fertile amphiploid population. Further selection was made for improved fertil- Slender wheatgrass (fig. 10) is a diverse complex of ity, seed production, seed size and vigor, herbage several taxa. It is a native, short-lived, cool-season, production, leafiness, and drought tolerance (Asay strongly tufted perennial that occasionally produces and others 1991). rhizomes (Cronquist and others 1977; Goodrich and Seed set of the SL-1 population approaches that of Neese 1986; Hitchcock 1951; Welsh and others 1987). the parental species, but seed fertility varies between Stems are green or bluish green and 20 to 60 inches 34 and 50 percent (Maughan 1988). The hybrid differs (0.5 to 1.5 m) tall. Sheaths are smooth and glabrous or from both parents in being leafier and producing rarely hairy. Blades are ascending to lax. They are larger culms and leaves. Rhizome development ranges usually narrow and flat or involute, 2 to 10 inches (5 to from a caespitose growth habit to plants that spread 1 to 25 cm) long, 0.08 to 0.16 inch (2 to 6 mm) wide, and 3.3 ft (0.3 to 1.0 m) per season (Asay and others 1991). scabrous to pilose on the upper surface. Spikes are 1.6 SL-1 exhibits usefulness in plantings on sites where to 8 inches (4 to 20 cm) long and stiffly erect. Spikelets either parent occurs. The SL-1 germplasm is quite are overlapping, 0.35 to 0.60 inch (9 to 16 mm) long, and vigorous and productive, supplying forage and re- three to seven flowered. Glumes are glabrous or maining green late into the summer. Considerable fall scaberulous, 0.25 to 0.60 inch (6 to 15 mm) long, 0.04 to greenup occurs, extending the period of grazing. The 0.12 inch (1 to 3 mm) wide, and abruptly tapered to an plant is also useful as a ground cover and could be acute or awn-tipped apex. Lemmas are blunt or end in planted to reduce erosion in semiarid areas. Plants a short awn. The second glume is about two-thirds the establish well by direct seeding and appear capable of length of the lemma. controlling weeds. Slender wheatgrass and four related taxa described by Hitchcock (1950) have been combined as varieties of

342 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

characteristic. Long-term ecological studies on the Wasatch Plateau of central Utah revealed that this species was one of the few native grasses to persist, although at a low density, after many years of heavy grazing (Monsen and others 1996). As grazing was discontinued, this species regained dominance rather quickly in some areas, but its position fluctuated with changes in climatic events and recovery of associated species. Slender wheatgrass has been able to persist in many overgrazed mountain herblands. It persists on sites where annual weeds, including tarweed, have become dominant. It invades openings and spreads to occupy harsh sites. It has also been able to persist amid disturbances where gophers are extremely destruc- Figure 10—Slender wheatgrass grows at higher elevations than other native bunchgrasses and tive. It is not a weedy invader, but it is able to persist is often used for reseeding forest disturbances and spread due to its seeding abilities and short-term (RMRS photo). growth characteristics. Plant Culture

Agropyron trachycaulum or placed in synonymy with Slender wheatgrass produces an abundance of seeds this species (Cronquist and others 1977). Arnow (1987) when grown on wildland sites or in cultivated fields. followed Gould (1947) in treating the complex in Hafenrichter and others (1968) reported that seed Elymus. Bearded wheatgrass is the most important production of irrigated plantings averaged 800 pounds taxon included in this group. It is similar to slender per acre (896 kg per ha). Plants are self-fertile wheatgrass in areas of occurrence and growth (Fulbright and others 1982), and viable seed crops are characteristics, but it is more productive, robust, and produced most every year. Seeds are large, detach longer lived (Hafenrichter and others 1968). easily, and can be combined and cleaned without difficulty. Seeds retained in warehouse storage lose Ecological Relations and Distribution little viability after 5 to 10 years. Slender wheatgrass normally requires cool Slender wheatgrass occurs in Alaska, much of prechilling for germination. Fall seeding is recom- Canada, and southward through the West, Central, mended, but seeding high mountain ranges in early and Northeastern portions of the United States, and in spring or summer is successful if summer rains main- Northern Mexico (Arnow 1987). It requires at least 15 tain moist seedbeds. Stewart and others (1939) recom- to 20 inches (38 to 51 cm) of annual precipitation. mend seeding 5 to 12 pounds of seed per acre (5.6 to Consequently, it is usually found at higher elevations 13.4 kg per ha). Seeds benefit from drill seeding and and on deeper soils than most other wheatgrasses. In placement of seeds in the soil at depths of 0.5 to 1.0 the Midwest, it is largely limited to soils with moder- inch (1.3 to 2.5 cm). This species can be seeded in ate water-holding capacity (Sarvis 1941). Slender mixtures with other grasses and broadleaf herbs. It is wheatgrass normally grows in association with aspen, an important native grass that can be seeded with spruce-fir, and mountain herblands (Mueggler and slower developing herbs without suppressing their Campbell 1986). It is not tolerant of dense shade, and seedling development. becomes quite abundant following timber cutting and Slender wheatgrass can serve as an important pio- fires. It is a common species in open park herblands neer species; its seedlings are vigorous and capable of (Eckert 1975; Ellison 1954) and open areas in moun- establishing on harsh sites. In addition, it is able to tain brush and ponderosa pine forests. It is less impor- establish and compete with weedy species, particu- tant in pinyon-juniper woodlands, but it can be quite larly cluster tarweed and knotweed. It can also be abundant in local areas. Slender wheatgrass occupies interseeded into sites disturbed by grazing that sup- some mountain big sagebrush benchlands, and it is a port species such as western yarrow, Letterman common constituent in river bottoms and on alluvial needlegrass, dandelion, and Rydberg penstemon to soils in stream and valley bottoms. improve diversity and stimulate natural recovery. Slender wheatgrass can be a relatively short-lived Because it is short lived, the density of slender wheat- species, but it reseeds and spreads well by natural grass in undisturbed herblands fluctuates annually, seeding, exceeding most other wheatgrasses in this recovering through natural seeding.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 343 Chapter 18 Grasses

Uses and Management planted in forested regions following fires to provide soil protection. New seedings establish quickly, fur- Slender wheatgrass was one of the first native grasses nishing excellent cover the first growing season. Plants included in early restoration plantings (Sampson 1913). establish well on open, exposed soil. This grass is also It became a widely used species for planting range and seeded on overgrazed ranges where soil stability is a wildland sites in the Intermountain West (Forsling concern (Plummer and others 1955). Price (1938) noted and Dayton 1931), the Northern Great Plains, and that differences occurred among ecotypes, and found Canada (Sarvis 1941). It was initially recognized as a that successful plantings could be established in low- desirable forage species that is easily established and elevation communities if adapted ecotypes were adapted to a wide range of sites (Forsling and Dayton seeded. Bridges (1942) reported the success of seed- 1931). Slender wheatgrass was not used in cultivated ing abandoned farms, dry meadows, and mountain pastures until specific cultivars were developed for mesas in New Mexico, using local ecotypes of slender that purpose. The short-lived nature of this plant wheatgrass. resulted in the unexpected decline of some early Slender wheatgrass provides a useful native substi- seedings, but as this characteristic was better under- tute for intermediate wheatgrass, smooth brome, and stood, slender wheatgrass became an important reveg- fairway crested wheatgrass seedings in areas where it etation species. is adapted. It can be used in mixtures with other Slender wheatgrass is a dual-purpose species. natives to reestablish native communities. Its wide Growth begins in midspring and continues through range of adaptation from mountain big sagebrush summer and fall if moisture is available. It is usually communities (Beetle 1952) to subalpine types (Stewart more preferred by cattle than by sheep (Vallentine and others 1939) makes this species a useful plant for 1985). Its foliage cures well and provides useful winter restoring native understories and rehabilitating harsh herbage (Sarvis 1941). Although slender wheatgrass disturbances. produces a moderate amount of herbage, it is highly nutritious and palatable. However, the plant can be- Varieties and Ecotypes come “stemmy” as it reaches maturity. Slender wheatgrass withstands intense use, but it is not as tolerant Five slender wheatgrass cultivars have been re- of grazing as western wheatgrass. Plants remain vig- leased. Because this species includes numerous re- orous with moderate to heavy grazing. Grazing at the gional ecotypes, it is important that adapted sources time of flowering and seed set, however, can diminish are planted. Some seed can be harvested from wild- plant vigor. Slender wheatgrass is preferred by elk, land stands, but most seed must be field grown to bighorn sheep, deer, and other wildlife (Hallsten and supply the amounts required for larger projects. others 1987). It provides forage and cover for wildlife. ‘Adanac’ slender wheatgrass was released in 1990 Animals seek and graze the green herbage throughout by Agriculture Canada for use in hay and pasture the summer. production in Saskatchewan. It is slightly more pro- Native and seeded stands of slender wheatgrass are ductive than ‘Revenue’ for hay production. It is not often managed as meadow hayfields or as pastures. commonly grown in the Western United States. New plantings grown under cultivation remain highly ‘Primar’ slender wheatgrass was cooperately reproductive for 3 or 4 years; yields diminish substan- leased by the Washington, Idaho, and Oregon Agricul- tially thereafter. The cultivar ‘Primar’ was developed tural Experiment Stations; the National Resources for pasture use and is seeded with sweetclover or red Conservation Service Plant Materials Center at Pull- clover in conservation and pasture plantings man, WA, and the Agricultural Research Service, (Hafenrichter and others 1968). Field production of Plant Science Research Division in 1946. It was devel- mixed seedings can be maintained at a higher level for oped from materials collected in 1933 near Beebe, MT. longer periods of time than if the grass is seeded alone. Primar is a vigorous, early-growing, and long-lived Slender wheatgrass is used primarily to restore selection. It is primarily seeded with sweet clover in disturbances and rehabilitate native communities. It pasture mixtures, conservation seedings, or as a green is widely planted to reestablish desirable cover on manure crop to improve soils (Alderson and Sharp mountain meadows and open park herblands. It can 1994). Early spring growth exceeds that of other col- be interseeded into existing cover with minimal site lections, and plants remain green throughout the preparation. Once plants become established and seed summer. Plants generally persist for fewer than 5 crops develop, new plants establish from seed. Native years on sites receiving less than 15 inches (380 mm) stands can also be managed to allow for natural of annual precipitation; they are much longer lived on recovery. Slender wheatgrass is commonly seeded in sites receiving more than 18 inches (460 mm) of an- mixtures with other grasses and broadleaf herbs. nual rainfall (Schwendiman and Law 1946). Seedings Slender wheatgrass can be used to quickly restore are extremely vigorous. When grown for hay, this ground cover and control erosion. This species can be cultivar provides abundant yields. Plants are free of

344 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses smut, and the species can be seeded for pasture use on Hitchcock and others (1969) applied the name Agrostis a wide range of soils. alba and recognized three varieties: A. a. var. alba, A. The Natural Resource Conservation Service Plant a. var. palustris, and A. alba var. stolonifera. These Materials Center at Bridger, MT, and the Montana varieties had previously been treated at the species and Wyoming Agricultural Experiment Stations re- level (Hitchcock 1950). Other more current treat- leased ‘Pryor’ in 1988. It was collected along a dry ments, including Arnow (1987) and Cronquist and drainage in a saline upland between Pryor and the others (1977), combined these varieties into A. Beartooth Mountains of Montana. It produces large stolonifera. seeds and vigorous seedlings. Pryor has been used Redtop, or carpet bentgrass, is an extremely long- primarily for conservation and reclamation planting lived rhizomatous and often-stoloniferous perennial in Montana and Wyoming (Alderson and Sharp 1994). that sometimes forms large mats (fig. 11). The pres- It is drought tolerant and adapted to saline soils. ence or absence of stolons, rhizomes, or both is re- Revenue slender wheatgrass was developed and ported to be habitat related. Stems are erect to decum- released in 1970 by Agriculture Canada, Saskatoon, bent and range from 0.7 to 4.9 ft (0.2 to 1.5 m) in height. Saskatchewan, from seed of a single plant collected in Leaf blades are flat or folded, smooth to strongly 1963 near Revenue, Saskatchewan. It is recommended roughened, and 0.08 to 0.8 inch (2 to 20 mm) wide. for use as a pasture variety because stands can be Ligules on the upper leaves are 0.2 to 0.28 inch (5 to 7 maintained for 3 to 5 years. This cultivar is superior to mm) long. Panicles are oblong to elliptical, 1.6 to 15.7 Primar in salinity tolerance, establishment, forage inches (4 to 40 cm) long, and 0.4 to 5.9 inches (1 to 15 production, and seed yields. It provides considerable cm) wide with the branches eventually spreading. leafy herbage, and plants are free of smut (Alderson Spikelets are single flowered, 0.06 to 0.1 inch (1.5 to and Sharp 1994). Revenue is widely seeded in pas- 3.5 mm) long, and green to reddish purple with an tures and on rangelands in the Western United States. awnless, or rarely short-awned lemma. The common ‘San Luis’ was cooperatively released in 1984 by the name redtop refers to the characteristic reddish hue of Colorado, Utah, and New Mexico Agricultural Experi- the panicle. ment Stations; the National Resources Conservation Service; and the Upper Colorado Environmental Plant Ecological Relationships and Distribution Center at Meeker, CO. It was developed from plants collected in San Luis Valley, Rio Grande County, CO. Redtop is one of the most important and widely Plants establish quickly, provide effective cover, and distributed members of its genus in the United States. are relatively long lived. San Luis is recommended for Native to Eurasia and North Africa, it was introduced sites above 5,800 ft (1,800 m) that receive at least 14 to North America prior to 1750 as a lawn, meadow, and inches (350 mm) of annual precipitation (Alderson and pasture grass. It now occurs throughout the northern Sharp 1994). It is widely seeded throughout the temperate region, growing in shallow water, wet mead- Intermountain area, particularly on drier sites than ows, and along streambanks on moderately moist are seeded with Revenue. It is commonly seeded on range and wildland sites, including roadway and mine disturbances.

Agrostis stolonifera Redtop or Carpet Bentgrass ______Synonyms Agrostis alba Agrostis palustris Agrostis gigantea Agrostis depressa

Description Figure 11—Redtop, a long-lived, mat-form- The taxonomic history and resulting synonymy of ing perennial introduced as a lawn, hay, and pasture grass, has spread widely and now this species is complex (Kartesz and Kartesz 1980). occurs in moist areas of many Western plant Hybridization has further confused the issue communities (photo courtesy of John Kinney, (Bradshaw 1958). Linnaeus based A. alba on a Poa, USDA Forest Service, Rocky Mountain Re- apparently P. nemoralis (Cronquist and others 1977). search Station, Boise, ID).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 345 Chapter 18 Grasses pinyon-juniper, aspen, and spruce-fir communities does grow through the summer if moisture is avail- (Arnow 1987; Cronquist and others 1977; Hitchcock able, and it can provide green herbage when foliage of 1950; Hitchcock and others 1969). The species occurs other species has dried. Redtop is not highly palatable, over a broad elevation range, from sea level to about and it does not tolerate close and repeated grazing. 10,000 ft (3,050 m), but mostly below 8,000 ft (2,440 m) However, it is useful for game and livestock. (USDA Forest Service 1937). Redtop receives considerable use in some areas. It is seeded in wet and semiwet meadows in aspen and Plant Culture mountain herblands, particularly in areas where erosion is a major concern (Stubbendieck and Jones Plants flower from June to early September. Abun- 1996). Redtop is also well adapted to acidic soils and dant seed crops are produced most years. Seeds are has promise for treating disturbances at high eleva- easily harvested, but they shatter readily when ma- tions. It is a vigorous grower and will form good turf in ture. Seed production fields yield about 75 pounds per a short time, making it valuable for use as a soil binder acre (84 kg per ha) (De France 1953). Viability of newly in reclaiming eroded drainages and stabilizing slopes, harvested seed is generally high. Seedlots are nor- banks, gullies, and waterways. It is also commonly mally cleaned to at least 90 percent purity; germina- seeded in mixtures with other species, including al- tion is often at least 90 percent. There are 5 to 6 million falfa, to provide pasture and hay (Weintraub 1953). seeds per pound (5.6 to 7.2 million per kg) (De France Redtop and several other stoloniferous species intro- 1953). duced from Europe are used extensively in turf cul- Redtop seeds are small and benefit from surface ture, especially for use on golf courses. seeding. Planting depth should not exceed 0.25 inch Redtop has been seeded to protect disturbed ripar- (0.63 cm). The species establishes well by broadcast ian sites and high-elevation herblands because seed of seeding followed by light harrowing to cover the seed. many desirable native species is not available. This technique works especially well when spring or Plantings have provided excellent ground cover and fall seeding on wet surfaces, which may crust if com- herbage, but they have maintained dominance on pacted by drill seeding. Seeding rates can be easily planted sites. Natural recruitment of native species adjusted to provide the desired seed density. A carrier has not occurred. Redtop should not be recommended may be added to regulate the seeding rate and seed or seeded if recovery of native species is desired. distribution. Redtop is normally not seeded alone, particularly in irrigated or dryland pastures. When Varieties and Ecotypes planted in mixtures, seeding rates vary between 2 and 4 pounds per acre (2.2 and 4.5 kg per ha). Rangeland ‘Streaker’ was developed in Idaho for pastures, rec- and watershed seedings normally do not require lamation, and turf plantings. It is presently the only heavier seeding rates. cultivar available (Jacklin and others 1989). Although redtop seeds germinate early, seedlings are weak and can be suppressed by other species. Thus, establishment is often poor. Mature stands, Alopecurus arundinaceus however, are very persistent. Plantings established Creeping Foxtail over 50 years ago on the Wasatch Plateau have sur- Reed Foxtail ______vived and flourished. Plants develop a low growth form and spread by producing a vigorous sod. Redtop Synonym does not spread aggressively beyond the original planting site in rangeland seedings, irrigated pastures, or Alopecurus ventricosus seed production fields. It does increase in density, and spreads by extension of the root system. Alopecurus pratensis Uses and Management Meadow Foxtail ______Redtop is recognized as a useful cultivated grass for Synonym wetland areas. It has been used extensively with other None. species in irrigated pastures (Weintraub 1953). Its area of distribution is similar to that of Kentucky Description bluegrass, but it is more tolerant of acidic, poorly drained, and clay soils (Balasko and others 1995). It The genus Alopecurus consists of about 50 species responds well to fertilization, but it is not a high- distributed throughout the Temperate and Arctic Zones producing species for hay production. Although other of the Northern Hemisphere (Boe and Delaney 1996). wet meadow species may be more productive, redtop Its name, Alopecurus, is derived form the Greek words

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Alopex meaning fox, and oura meaning tail, referring to the characteristic cylindrical panicle. The two most common forage species in this genus are meadow foxtail and creeping or reed foxtail. These two perennial species are similar in taxonomy, growth form, site requirements, and uses. Consequently, they will be discussed together. Linnaeus named meadow foxtail A. pratensis in 1753, and the name has never been changed. Hitchcock (1950) classified creeping foxtail as A. arundinaceus, and this name continues in use. Creeping foxtail and meadow foxtail are separated taxonomically by the position and extension of the awn. The awn of creeping foxtail arises below, or rarely above, the middle of the lemma. It is straight or slightly bent, and measures 0.04 to 0.1 inch (1 to 3 mm) in length. The awn is either included or exerted (Arnow 1987). The awn of meadow foxtail is attached near the base of the lemma and is bent or geniculate. Creeping foxtail is a rhizomatous perennial that ranges from 1.6 to 3.9 ft (0.5 to 1.2 m) in height (fig. 12). Culms are solitary or in small clusters. Leaf blades are flat, 0.3 to 0.5 inch (8 to 12 mm) wide, glabrous above and rough below. Ligules are 0.04 to 0.20 inch (1 to 5 mm) long. Panicles are purple or dark colored, spikelike, cylindric, 1.6 to 3.9 inches (4 to 10 cm) long, and up to 0.3 inch (8 mm) wide. Spikelets are one flowered and laterally compressed, disarticulating below the equal glumes. Diagnostic characters for this species include the glume tips, dark purple spikelets, and shortly exerted awns (Arnow 1987). Figure 12—Creeping foxtail, a rhizomatous in- Meadow foxtail is a stout, nonrhizomatous peren- troduction, grows on wet sites over a wide nial ranging from 1.3 to 2.6 ft (4 to 8 dm) in height, but elevational range. Seeded to provide hay, for- sometimes reaching 3.3 ft (1 m) (fig. 13). Roots some- age, and erosion control, it has become weedy times develop from the lower culm nodes. Leaf blades along waterways (photo courtesy of Kevin Jensen, USDA Agricultural Research Service, are flat, roughened on both surfaces, and 0.12 to 0.24 Logan, UT). inch (3 to 6 mm) wide. Ligules are 0.08 to 0.14 inch (2 to 3.5 mm) long and covered with short, fine, soft hairs. Ligule margins are irregularly serrate to entire. Meadow foxtail was introduced in North America in Panicles are 1.4 to 3.4 inches (3.5 to 8.5 cm) long and the mid-1800s (Wheeler 1950), and it has naturalized 0.2 to 0.3 inch (5 to 7.5 mm) wide. Spikelets are ovoid across Canada and the northern half of the continental and one flowered. Glumes are subequal, often with United States (Hitchcock 1950; Scoggan 1978). Both green nerves, and relatively large, measuring 0.12 to species are long lived, winter hardy, and adapted to 0.20 inch (3 to 5.2 mm) in length. Keels of the glumes high-elevation meadow sites, including those with have long, soft, unmatted hairs; those on the lateral poorly drained, acidic soils. Both have a high tolerance nerves are appressed. Lemmas are 0.12 to 0.20 inch (3 to flooded and frozen soils (Stroh and others 1978) and to 5 mm) long and usually glabrous, but sometimes are capable of persisting on sites where standing with hairs near the keel apex. water may be present for more than a month. They are seeded as pasture and hay crops in wet meadows and Ecological Relationships and Distribution have spread along waterways and moist drainages. Creeping foxtail and meadow foxtail are widespread in Europe and temperate Asia (Strelkova 1938). Creep- Plant Culture ing foxtail was introduced in the United States in 1935 Panicles mature from the apex toward the base. (Stevens 1963), and is now naturalized in the Great Seeds near the panicle apex may shatter before seeds Plains, Pacific Northwest, and Intermountain regions near the base mature, and seeds can shatter within a (Hafenrichter and others 1968; Sutherland 1986).

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furrows at a depth of less than 0.5 inch (1.3 cm). Surface-sown seed should be harrowed or covered using a light drag. The planted unit consists of a caryopsis enclosed in a light, fluffy, membranous covering that can bridge or flow irregularly in most conventional seed boxes. Consequentially, adaptors are required to move seed uniformly through drills or other planting machinery. Creeping foxtail has been seeded primarily on range and wildland sites. It has established well without irrigation from both drill and broadcast seedings. Dispersed seeds germinate quickly, allowing plants to spread and develop competitive stands rather quickly. Mature plants compete well with other species, and ultimately dominate areas where the species is adapted. Although seedlings of creeping foxtail and meadow foxtail appear to be weakly competitive when seeded in cultivated pastures with other species, both plants are aggressive natural spreaders on rangelands (Plummer and others 1968). They spread by root proliferation and natural seeding, particularly the latter (Lewis 1958b).

Figure 13—Introduced in the mid 1800s, Uses and Management meadow foxtail has become naturalized on high-elevation sites in Canada and the North- The foxtails were among the first perennial grasses ern United States (RMRS photo). used to rehabilitate disturbances on mountain rangelands. Both species are well adapted to subalpine grasslands, aspen forests, and mountain brushlands. They have been widely planted in irrigated and few hours after reaching maturity (Wheeler and Hill nonirrigated pastures in mountain valleys, wet and 1957). Consequently, seeds must be harvested when semiwet meadows, and riparian zones. They have also the greatest number of seeds are mature, but before performed well on drier sites where supplemental they disperse. Seeds tend to darken or turn black irrigation is not available. Plants are equally adapted during maturation, a trait that aids in selecting har- to meadow sites in lowlands or in subalpine sites at vest dates. Seeds can be combine harvested by raising elevations over 10,000 ft (2,857 m). Both foxtails are the cutter bar above the leaves to remove seed heads shade tolerant, and have persisted well as understory and some stem material. Harvesting can also be ac- species in aspen forests. Regrowth following grazing complished using strippers that dislodge the seeds by can occur beneath an aspen canopy throughout the flailing or brushing. Seed can be harvested when summer, even though these sites are shaded (Piper slightly green if it is allowed to dry before processing. 1944). Seed yields are not usually as high as for other pasture Foxtails are usually planted as pasture or forage species, but yields between 268 to 400 pounds per acre species. They begin growth in early spring, earlier (300 to 450 kg per ha) have been reported (Hafenrichter than most other forage species, including timothy and 1968). Meadow foxtail and creeping foxtail usually orchardgrass. Plants often emerge through snow cover produce good-quality seeds that can be cleaned to and begin growth when the soil is still frozen (Stroh relatively high standards. Seed germination and pu- and others 1978). In addition, they produce an early rity each normally exceed 80 to 85 percent. seed crop that is used by a variety of animals. Plants Spring or fall seedings can be successful in wet continue growth during mild winters (Hafenrichter meadows, as seeds germinate rapidly on moist seed- and others 1949). beds. Ten to 14 pounds of seed per acre (11.2 to 15.7 kg Geese, big game animals, and other wildlife species per ha) is adequate for planting irrigated pastures. In are attracted to the foxtails throughout the growing most rangeland situations, lower rates provide ad- season, but particularly in spring and fall (Boe and equate stands. When seeded in mixtures, particularly Delaney 1996). Plants produce considerable herbage with legumes or other pasture grasses, 3 to 5 pounds early in the season, and recover quickly from grazing per acre (3.4 to 5.6 kg per ha) is sufficient. Seeds can or clipping, as only about 20 percent of all shoots are be distributed on the soil surface or planted in shallow reproductive (Rumburg and Siemer 1976). Domestic

348 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses livestock are also attracted to foxtail plantings in survives periods of complete inundation and exhibits midsummer because new shoots are produced through- moderate tolerance of saline and alkaline soils. out the growing season. Upland game birds seek the cover and seeds provided by both species. Both foxtails are best used to support high stocking Aristida purpurea management, but have reportedly depressed animal Purple Threeawn or gains, possibly due to the presence of an unidentified Red Threeawn ______antiquality component (Rode and Pringle 1986; Voigt and Sharp 1995). Both species are grown for hay Synonyms production, particularly in high-elevation areas with short, cool growing seasons where few hay or pasture Aristida fendleriana species are adapted. On such sites early summer Aristida glauca pastures are important, particularly hayfields that Aristida wrightii can be cut or harvested early, but which can recover Aristida longiseta and be intensively grazed later in the season. Foxtails are commonly seeded with legumes to im- Description prove yields and forage quality (Boe and Delaney The perennial Aristida complex has been treated as 1996). They have been seeded with various clovers, five separate species as listed above. The morphologi- birdsfoot trefoil, and alfalfa for pasture and hay pro- cal characters used to distinguish the species involved duction. On rangelands they are more commonly seeded a continuum of features including beak length (awn with alfalfa, although some plantings with cicer column), beak twisting, awn length, panicle length, milkvetch have also produced excellent pastures. panicle branching, lemma roughness, and leaf posi- Creeping foxtail plants have been noted to spread as tion (Cronquist and others 1977). These characters much as 4 ft (1.2 m) over a 3-year period (Stroh and apparently vary from one extreme to another in any others 1978). Consequently, they can outcompete as- given population. Some even appear to differ on the sociated species, including sedges and rushes. Plants same plant from year to year depending upon chang- also form dense rooting systems that provide excellent ing environmental conditions. Considerable differ- soil protection. Hart and others (1980) reported that ences in plant stature and leaf and stem numbers have creeping foxtail suppressed Canada thistle. These also been noted. Due to this extensive variability, species have been used on high-elevation disturbances Cronquist and others (1977) included all taxa in to stabilize watersheds, waterways, streambanks, and Aristida purpurea and recognized three varieties: aspen disturbances. In these situations, however, they purpurea, glauca, and robusta. Regional ecotypes oc- have gained dominance and prevented the recovery of curring in specific environments have also been de- native species. scribed (USDA Forest Service 1937). Plants of the threeawn complex are tufted perenni- Varieties and Ecotypes als that range from 0.5 to 2.1 ft (1.5 to 6.5 dm) tall. ‘Dan’ meadow foxtail was developed from material Culms are smooth and hairless and sometimes collected in Poland (Alderson and Sharp 1994). It is branched at the lower nodes. There are long, soft, used primarily on low-elevation sites, while ‘Moun- white hairs at the apex of the sheath. Leaf blades are tain’, developed in Ontario, Canada, is a more upland inrolled or involute, and rarely flat, glabrous to slightly variety that is seeded in semiwet meadows associated roughened on the upper side, 0.04 to 0.08 inch (1 to 2 with mountain brush, aspen, and conifer communi- mm) wide, and 2 to 3.9 inches (5 to 10 cm) long. They ties. Mountain greens up in early spring and again in are often curved or flexuous on small plants and fall. Neither variety has been seeded to any extent in straight on large ones. Ligules are scarcely 0.02 inch the Intermountain West. (0.5 mm) long and consist primarily of a fringe of hairs. ‘Garrison’ creeping foxtail is a pasture and hay grass Panicles are 2.4 to 9.8 inches (6 to 25 cm) long and that is adapted to waterways, wetlands, and other narrow to somewhat spreading. Panicle branches are areas with high water tables. It was developed from usually appressed and closely clustered. Spikelets Western European material growing near Max, have narrow, linear glumes of uneven lengths that are McLean County, ND (Alderson and Sharp 1994). Gar- sometimes awned. The lemma is about 0.4 to 0.6 inch rison is seeded extensively in wet and semiwet pasture (9 to 15 mm) long (including the awn column), often renovations. It can be seeded directly into weak stands grading into a thin, twisted awn column or beak. The of forbs and perennial grasses, including saltgrass. Its three awns that radiate from the awn column are seedlings establish slowly, but established plants are subequal in length and have a purple hue. Plants highly productive and spread aggressively. Garrison flower from March to September (Cronquist and others 1977).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 349 Chapter 18 Grasses

Ecological Relationships and Distribution heavily grazed nor eliminated as quickly by grazing as bluebunch wheatgrass. Purple threeawn is commonly found in dry, sandy soils of desert valleys, plains, mesas, and foothills. It is Plant Culture distributed from British Columbia throughout the Western and most of the Central States and south into Plants begin growth in early spring, generally by Central Mexico at elevations ranging from 2,500 to April (Evans and Tisdale 1972). Flowering occurs in 10,000 ft (763 to 3,050 m) (fig. 14). In the United States mid-June and seeds mature in early September. Some it is especially important in the Southwest. Purple viable seeds are normally produced, even during dry threeawn is often abundant along stock driveways, years (USDA Forest Service 1937). The divergent roads, and abandoned farmlands. It invades distur- awns and sharp callus are important for natural bances where animals have burrowed, or where plow- dispersal and seeding, as they anchor and position the ing or grazing have reduced or eliminated native seed in the soil (Evans 1967). vegetation. It is often found on south- and west-facing Purple threeawn is not widely used in revegetation slopes where soil temperatures may be abnormally projects. At present, most seed is collected from wild- high (Evans 1967). At midelevations it grows with land stands using modified hand or vehicle-mounted sand dropseed, needle-and-thread, and bluebunch equipment. Seed collection and processing is hindered wheatgrass. At lower and warmer sites it occurs with by the long awns. Removal of the awns is necessary to sideoats grama, Indian ricegrass, and bottlebush reduce volume for storage and to plant seeds with squirreltail. conventional drills or other seeding equipment. Fall Purple threeawn often grows in nearly pure stands. seeding at a rate of 6 to 10 pounds per acre (6.7 to 11.2 It may also be found intermixed with other native kg per ha) is recommended if the species is planted bunchgrasses as a codominant species or as a minor alone. Drill seeding is generally required to place the component of bunchgrass/sagebrush communities. seed at a depth of 0.5 to 1.0 inch (1.3 to 2.5 cm). Evans Daubenmire (1970) suggested that purple threeawn and Tisdale (1972) reported purple threeawn seeds distribution patterns indicate that its presence and required treatment at high temperature to initiate distribution are regulated by edaphic conditions. Purple germination. Little seed germination was recorded threeawn is reported to invade disturbances previ- until seeds were subjected to air temperatures exceed- ously occupied by bluebunch wheatgrass, and it is ing 104 ∞F (40 ∞C). often considered an early seral species (Evans 1967). Purple threeawn has been successfully seeded in However, this species is widespread and often present mixtures with other native species. It competes well as in undisturbed situations (Küchler 1964). It is not as a seedling. Root systems develop rapidly, allowing young plants to establish and persist, even during periods of moderate drought. Plants may require 3 to 5 years to fully develop. They recover well following disturbance, spreading from natural seeding. Devel- oping stands and mature stands allow reestablishment of associated native shrubs and forbs. Purple threeawn recovers quickly following burning due, in part, to the production of some seed each year. Seeds tolerate high temperature resulting from fires (Sampson 1944), and some may persist in the soil as a seed bank, aiding in natural recovery over time. Nu- merous small seedlings of purple threeawn may be found following fire and other disturbances. This plant is especially adapted to rapidly establish and spread due to the awns that facilitate dispersal and planting. Small and developing seedlings are able to compete with cheatgrass and other weeds. Seedling survival and growth of young plants in weedy competition indicates this species could be used to contain weeds and facilitate natural community recovery. Mature purple threeawn plants respond somewhat differently to burning. Fire can kill clumps that have an accumulation of large amounts of litter, but in most situations Figure 14—Purple threeawn is distributed from Brit- burning may remove dead material but few plants are ish Columbia to Mexico (RMRS photo). killed (Evans 1967).

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Uses and Management Description Although not generally considered a highly palat- Linneaus (1753) described tall oatgrass (Arrhena- able species, purple threeawn produces considerable therum elatius), naming it Avena elatior. Its current early spring forage that is used by livestock and big binomial, Arrhenatherum elatius, was assigned by game animals (Dittberner and Olson 1983). The USDA Presl (1819). Tall oatgrass is a short-lived perennial Forest Service (1937) identified it as an important bunchgrass, often rooting from the first nodes. The forage plant in some areas, but noted that consider- plant is erect and not truly rhizomatous. Culms arise able differences existed among ecotypes. As awns from the bulblike base and are loosely tufted, ranging develop, plants become less attractive to grazing ani- from 2.3 to 5.9 ft (7 to 18 dm) in height. Leaf blades are mals, and can cause injuries if the awn attaches to or flat, smooth to somewhat roughened, and 0.1 to 0.2 penetrates the skin, face, and eyes. Palatability is inch (3 to 6 mm) wide. Ligules are 0.04 to 0.12 inch (1 rated as poor to fair as the awns appear. The nutri- to 3 mm) long, obtuse, and covered with very fine, tional value of the species becomes somewhat more ciliate hairs. Panicles are 5.5 to 9.8 inches (14 to 25 cm) important in midsummer, fall, and winter months long, narrow, and bear flowers to the base. Spikelets (Evans 1967), as it maintains some green foliage when are shiny and two flowered with the first floret stami- other species are normally dry. nate and the second perfect. The florets disarticulate Purple threeawn is an important understory species as a unit. Glumes are membranous and unequal, in many shrubland communities that occupy dry slopes measuring about 0.2 to 0.3 inch (5 to 7 mm) in length. and sites critical as winter and spring-fall ranges for The awn of the first lemma is bent, about 0.4 to 0.6 inch big game. In addition, these sites are subject to inva- (10 to 14 mm) long, and it is attached midlength on the sion by annual weeds if the understory is disturbed. lemma. The awn of the second lemma is small, straight, The presence of annuals has resulted in a serious and attached at the lemma apex. It is rarely absent. decline of many woody species due to competition and Plants flower from late May through July (Arnow increased fire frequency. Consequently, it is impor- 1987). tant to maintain understory species that are compatible with associated shrubs. Ecological Relationships and Distribution Purple threeawn is adapted to disturbed sites and soil conditions where few other species exist. Kindell Tall oatgrass is native to Eurasia, but it is now and others (1996) reported that ecotypes differ in their circumboreal (Arnow 1987). Introduced to the United adaptation to local conditions. In many areas attempts States in 1807, it has been widely cultivated and is to replace these ecotypes with more palatable intro- now distributed across the United States (Weintraub duced perennial grasses have diminished species di- 1953) and Southern Canada. It is found in pastures versity and resulted in a loss of habitat. Restoring and and hayfields in the Pacific Northwest, the Inter- maintaining purple threeawn is essential to the health mountain and North-Central States, and the desert of many rangelands. Southwest. It is well adapted to subhumid and irri- Purple threeawn has been successfully seeded on big gated conditions (Hafenrichter and others 1949). Al- game ranges in central Utah, and on disturbed road- though primarily used as a pasture plant, it has ways and mined sites in the Intermountain West. It has escaped cultivation and spread to waste areas, par- established and developed when seeded in mixtures or ticularly those with rich soils. It has been seeded on alone. Seed prices remain high, as demand is sporadic. rangelands and watersheds in the Intermountain Seed production fields have not been developed. Use of region, and has spread to occupy sites in mountain this species will likely increase as reestablishment of brush, aspen, and mountain herbland communities. It native species on foothill and lower elevation sites tolerates shaded areas, and has spread to forested and becomes more important. wooded types. Plants are moderately salt tolerant, and have been planted in saltgrass meadows in the West Varieties and Ecotypes (Weintraub 1953) (fig. 15). There are no releases. Plant Culture Tall oatgrass is a rapidly developing species with Arrhenatherum elatius vigorous seedlings that is planted to provide pasture Tall Oatgrass ______and forage. It is also harvested for seed, as good seed crops are generally produced each year in cultivated Synonyms pastures and on most rangeland plantings. Seeds are borne on long, narrow panicles. Individual seeds are Avena elatior light, fluffy, and short awned. Seed crops can be Avena bulbosa

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seeded on areas such as mountain meadows characterized by cool, moist conditions. It is commonly seeded as a mixture with other species to provide diversity and improve forage quality and yields (Weintraub 1953). The hay is harvested early in the season, and it is palatable and nutritious (Irwin 1945). Tall oatgrass produces a reasonable amount of herbage in both pasture and rangeland plantings. Yields of few other species exceed those of this grass. Tall oatgrass is palatable and grazed by game and livestock. Its erect growth habit and abundant leaves provide considerable herbage in early spring. Exces- sive use in the early stages of growth can weaken the stand (Hafenrichter and others 1968). Although its palatability diminishes in midsummer and fall on rangeland sites (Plummer and others 1968), it cures well when harvested as hay (Stubbendieck and Jones 1996). Ordinarily, tall oatgrass begins to grow a few weeks earlier and remains green later in the fall than most other native grasses, thus extending the grazing period for livestock and wildlife (USDA Forest Service 1937). Plants retain only moderate amounts of edible forage in late fall and winter on game ranges. On Western rangelands, tall oatgrass has become an important species for seeding subalpine, aspen, and mountain brush communities. It is planted on range and watershed disturbances to quickly stabilize and Figure 15—Tall oatgrass has been widely protect the soil. It is also commonly used to seed forest planted on saltgrass sites across the West lands following wildfires. Plants are moderately salt (RMRS photo). tolerant, and the species has been used for seeding saltgrass meadows. Individual tall oatgrass plants are not long lived, surviving for 6 to 10 years. Natural recruitment occurs if associated vegetation does not fill in and occupy openings (Hull and Holmgren 1964). difficult to harvest because seeds shatter readily when Grazing management can affect long-term survival of mature. Light winds and combines or other machinery individual plants and seedings. can dislodge the seed and reduce the harvest. Seeds Although the ecological impacts of seeding tall are difficult to process. oatgrass on deteriorated aspen and mountain herbland Tall oatgrass can be difficult to seed through drills communities are not well understood, it is apparent because the seeds often clump together to form balls that seeding this species in disturbed situations does that impede the flow through the drill. This problem not seriously interfere with natural recovery of native can be corrected by processing the seed to separate the grasses and broadleaf herbs. Tall oatgrass was planted fluffy material. When seeded with seeds of other spe- on mountain herbland disturbances of the Wasatch cies, tall oatgrass seeds are often carried through the Plateau in the 1930s and 1940s. Excellent grass stands planting device in the mixture. Tall oatgrass seed can developed and soil erosion was stabilized. Within 10 to be distributed adequately by aerial or broadcast seed- 30 years tall oatgrass essentially disappeared. Natu- ing methods. Plummer and others (1968) reported ral recovery of native species began before the decline that seeds germinate without special treatment, and of the seeded grass. Obviously this grass can better uniform stands are easily established from either drill serve to provide initial ground cover and facilitate the or broadcast seedings. New plants grow rapidly and recovery of native species than more persistent peren- begin spreading from seed within 2 to 3 years if nial grasses, such as smooth brome, that are com- rodents or insects do not destroy the seed crops. monly used in these situations.

Uses and Management Varieties and Ecotypes Tall oatgrass has been used primarily as a pasture ‘Tualatin’ tall oatgrass was developed from materi- and hay crop in the Intermountain area. It is generally als collected in Chile, Italy, and Belgium (Alderson

352 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses and Sharp 1994) in an effort to provide a cultivar that does not shatter rapidly (Hanson 1972). It has been seeded at sites throughout the Intermountain West in Gambel oak thickets, aspen and conifer forests and associated openings, and in subalpine communities to provide soil stabilization, forage, and community enhancement. It is not aggressive and can be seeded with other perennial species. Tualatin grows well in full sunlight, and it also exhibits good shade tolerance. Plants are relatively short lived, but they will spread from seed if the seedheads are not grazed. Tualatin was selected for its rapid establishment and excellent forage characteristics. It has not been seeded to any extent in the Intermountain West. This cultivar is leafier than most common lines, but seed is not currently maintained or available.

Bouteloua curtipendula Sideoats Grama ______Synonyms Chloris curtipendula Cynodon curtipendula Andropogon curtipendula Melica curtipendula

Description Figure 16—Sideoats grama, a native warm season peren- Sideoats grama is a widespread, native, warm-sea- nial, is named for the spikes arranged largely along one side son, perennial midgrass. It is the largest grama spe- of the inflorescence (Hitchcock 1950). cies, reaching heights of 0.7 to 3.3 ft (0.2 to 1 m) (Stubbendieck and others 1986). Plants are tufted with scaly, slender to stout rhizomes. The species is descriptively named based on the characteristic spikes that are predominately on one side of the seedhead Ecological Relationships and Distribution (fig. 16). Culms are slender, erect, and smooth with a purplish tint at the nodes. Leaves are blue green and Sideoats grama is widely distributed throughout flat to involute. The ligule is reduced to a fringe of much of North America. It ranges from Saskatchewan hairs, approximately 0.04 inch (1 mm) long. Panicles east to Ontario, and from Michigan south to southeast- are 5.5 to 8.0 inches (14 to 20 cm) long with 25 to 80 ern California through Mexico to Central America to pendulous short-stalked, spikelike, branches that dis- Andean South America (Arnow 1987; Cronquist and articulate readily as a unit when mature. Each branch others 1977; Hitchcock 1950; Hitchcock and others produces two to eight spikelets, each with one perfect 1969; USDA Forest Service 1937), growing on plains, floret. There is sometimes a vestigial spikelet. Glumes prairies, and foothills. It often grows in canyons and on are papery and often purple tinged. The fertile lemma low-elevation mesas, scrublands, weedy meadows, open is three nerved with the lateral nerves ending as short woodlands, and steep rocky slopes at elevations from awns that are 0.12 to 0.24 inch (3 to 6 mm) long (Arnow 3,000 to 8,000 ft (915 to 2,440 m). It is especially 1987; Cronquist and others 1977; Hitchcock 1950; abundant in the central and southern mixed prairies Hitchcock and others 1969). There are two varieties of of North America. Rhizomatous B. c. var. curtipendula sideoats grama that are segregated by growth form: B. occurs over the vast majority of the species range, c. var. curtipendula arises from creeping rhizomes while nonrhizomatous B. c. var. caespitosa occurs in while B. c. var. caespitosa forms clumps. the Southwestern United States and in Central and South America.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 353 Chapter 18 Grasses

Sideoats grama is most abundant on fine-textured Because sideoats grama is naturally distributed soils ranging from deep to shallow, but it also occurs on over a wide area, ecotypes with different areas of sandy to dry, rocky soils (Nicholson and Bonham adaptation are abundant. Local seed sources or variet- 1977). It is better adapted to calcareous or moderately ies with known adaptation to the seeding area are alkaline conditions than to neutral or acidic soils. recommended for seeding. It is fairly drought resistant; root growth continues during dry periods (Weaver and Albertson 1956). Semi- Uses and Management nal roots rapidly extend deeper into the soil when subsurface soils are wet, reducing plant dependency Sideoats grama provides valuable forage for all on fluctuating moisture availability near the surface classes of livestock and big game (Dittberner and (Simanton and Jordan 1986). Abundant rhizome re- Olson 1983). Although plants are palatable and nutri- production may then occur following precipitation tious throughout most of the year, palatability is events (Weaver and Albertson 1956). greatest in summer and fall and declines in winter Sideoats grama is climax-dominant or codominant (Stubbendieck and others 1986; White 1986). This over much of its range (Great Plains Flora Association species differs from other warm-season species be- 1986). It also occurs in early seral communities on cause it produces important green herbage in spring. disturbed sites, and it generally increases following Mature stems are usually not eaten (Towne and drought (Weaver and Albertson 1956). In the Inter- Owensby 1983). Birds seek out the seed, and small mountain West, sideoats grama can be a climax indi- mammals feed on the seeds and foliage (Wasser 1982). cator in big sagebrush, pinyon-juniper, and ponderosa Sideoats grama has been successfully used in range- pine communities, and an associated species in shrub land seedings and on critically eroding areas, water- communities including black greasewood and true ways, and mined lands. If competitive species such as mountain mahogany. It is often found growing with pinyon, juniper, and big sagebrush are reduced, den- other grasses such as bluebunch wheatgrass, western sity as well as forage and seed production of sideoats wheatgrass, prairie junegrass, blue grama, and Idaho grama tend to increase. Following drought, sideoats fescue. grama generally increases dramatically. Although less tolerant of grazing than blue grama, sideoats grama Plant Culture tends to replace taller grasses on abused ranges. It will gradually decrease if continually grazed during the Sideoats grama reproduces from rhizomes, tillers, growing season. This species, however, has the ability and seed. Rhizome production and tillering are most to recover naturally from destructive grazing, but common in sandy soils. Natural reproduction from recovery can be slow in some situations. Strong natu- seed occurs when sufficient moisture is available for ral recovery is an important characteristic of this seed production, germination, and establishment species because many disturbances within its range (Simanton and Jordan 1986). Seed production and cannot be artificially seeded. Established seedings are quality varies with precipitation and grazing pressure fire tolerant. Sideoats grama is more site specific than (USDA Forest Service 1937). Germination is generally many other perennial grasses, and ecotypes should be rapid when sufficient moisture is available. Seedling planted on soils and sites where they are adapted. vigor is good compared to that of other warm-season Sideoats grama has proven only moderately adapted grasses, even on harsh sites. to harsh disturbances. It is receiving increased use as Seed can be cleaned and processed without damage. an ornamental grass. Seeds are small, but easily planted with most seeding equipment. Shallow planting at depths of 1 inch (2.5 Varieties and Ecotypes cm) or less is required. Seeds flow freely through drills, but they can be scattered by wind if broadcast aerially. ‘Killdeer’ was selected for its outstanding vigor, In more southern areas of the West, seedings con- leafiness, and fair seed production. It was originally ducted prior to summer rains are more successful. collected near Killdeer, ND, in an area that receives 15 Late fall and early spring seedings have been more inches (380 mm) of annual precipitation. It has been successful in the northern portion of its range. Drought seeded with good success in the Dakotas. tolerance and good germination and seedling vigor ‘Niner’ originated near Socorro, NM, at 4,200-ft contribute to its establishment and success. Because (1,280-m) elevation on a site that receives an average of its size and vigorous rate of growth, sideoats grama of 9 inches (230 mm) of annual precipitation. It dem- is quite competitive, and can be planted in mixtures or onstrates superior seedling vigor, strong establish- seeded alone. When seeded in mixtures with other ment, and high seed and forage production in dry species, it most often develops in small patches. It can areas. It is recommended for use in New Mexico, be successfully seeded with a number of native shrubs, Colorado, southern Utah, and northern Arizona. as it does not restrict seedling establishment.

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‘Pierre’ exhibits outstanding vigor, excellent leafy Arizona. It also occurs in southern California, Utah, forage production, and good disease resistance. It has and Wyoming. It is a climax species in a number of been successfully seeded in South Dakota and in the grassland communities and occurs in early seral com- surrounding States and Canadian Provinces. Its ori- munities on disturbed sites. It is most often found in gin is near Pierre, SD, in a 16-inch (410-mm) precipi- grasslands on dry, rocky hills and uplands and lower tation zone. elevation mesas on well-drained, sandy and gravelly ‘Vaughn’ originated near Vaughn, NM, in a 13.5- soils. Common associated species include dropseeds, inch (340-mm) precipitation zone at 5,930 ft (1,810 m). threeawns, broom snakeweed, mesquite, and yucca. It is best adapted to areas in western Texas, New Black grama can survive on harsh sites characterized Mexico, southern Colorado, southern Utah, and north- by low rainfall, high temperatures and evaporation, ern Arizona. It is a good seed and forage producer, and and strong winds (Nelson 1934). It is generally consid- it has received some use in the Intermountain West. ered tolerant of drought. During dry periods, however, large tufts have been observed to break up and form smaller clumps, with a reduction in ground cover, but Bouteloua eriopoda an increase in plant number. Black Grama ______Plant Culture Synonyms Stands are maintained and increased through seed Chondrosium eriopodum production, stolon proliferation from old tufts, and Bouteloua brevifolia tillering (Heizer and Hassell 1985). Reproduction varies widely with precipitation and grazing pressure Description (Canfield 1939). Two periods of flowering may occur in Black grama is an important native, warm-season, moist years (Canfield 1939). During a drought year, perennial grass that exhibits a variety of growth forms very little, if any, flowering occurs, and seed produc- over its range of occurrence. Tufted plants 10 to 28 tion is limited (USDA Forest Service 1937). In addi- inches (25 to 70 cm) in height develop from a hard, tion, uneven distribution of precipitation in summer knotty, woolly base. The slender culms are solid, may prevent seed from maturing (Nelson 1934). Black densely white-woolly, spreading, and sometimes grama seed fill and germination are often poor. Heizer stoloniferous at the lower nodes. They remain green up to the second node from the stem apex during some winters. Clusters of leaves emerge from the nodes in spring. Leaves are mostly basal, narrow, and pointed. They are generally twisted and involute near the top, hairy above, and smooth below. Roots are finely divided and generally confined to the upper 10 inches (25 cm) of soil (Arnow 1987; Cronquist and others 1977; Great Plains Flora Association 1986; Judd 1962). Inflorescences consist of one to six slender, comblike spikes borne on the sides of the inflorescence (fig. 17). The spikes are narrow, flaglike, and do not drop at maturity. Vegetative growth begins as soon as adequate moisture becomes available. The growth period usually lasts from July to October, depending upon temperature and precipitation. The vigor of black grama plants is largely determined by the amount of precipitation received during the previous summer (Canfield 1939). Flowering occurs from June or July through August. Seeds usually mature in late September and disperse in October and November (Nelson 1934).

Ecological Relationships and Distribution Figure 17—Black grama inflorescences consist Black grama is a major warm-season grass of arid of one to six slender, comblike spikes (Hitchcock and semiarid grasslands in Texas, New Mexico, and 1950).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 355 Chapter 18 Grasses and Hassell (1985) reported seed fills of only 7 to 10 ‘Nogal’ is less erect than ‘Sonora’ and adapted to percent for seed from native stands, while Campbell drier areas. Seed production is generally low, and and Bomberger (1934) reported that germination may supplies are limited and expensive (Alderson and be as high as 41 percent once every 10 years. Sharp 1994). In favorable years ground cover is increased through Sonora demonstrates outstanding leafiness, vigor, tillering, perhaps the most important means of regen- forage production, and vegetative spread, along with eration (Campbell and Bomberger 1934). On ungrazed good seed set and seed production. It is adapted across ranges, black grama forms uneven stands of large much of the species range. Seed is expensive and tufts that die in the center during drought, forming supplies are very limited. many smaller tufts. When favorable conditions pre- vail, these small tufts enlarge through lateral spread, and can again form a continuous cover (Nelson 1934). Bouteloua gracilis Stolons required two growing seasons to establish new Blue Grama ______plants. New stolons are produced during one growing season, while tillers develop and root during the sec- Synonyms ond growing season (USDA Forest Service 1937). Actinochloa gracilus Atheropogon gracilus Uses and Management Bouteloua oligostachya Black grama is used by all classes of livestock and wildlife (Stubbendieck and others 1986), and it was Description once a mainstay forage for livestock production in Blue grama is a low-growing, densely tufted, warm- many areas of the Southwest. It is highly palatable season perennial bunchgrass. The short rhizomes that and nutritious in summer and winter. During winter develop from its base often create thick mats. Culms and early spring it can provide forage when other are 6 to 12 inches (15 to 30 cm) tall. Leaves are grayish sources are scarce or unavailable (Canfield 1934). green and turn to gray or straw colored with age. They Although black grama can withstand recurrent graz- are persistent, numerous, mostly basal, and some- ing by livestock, it spreads very little on heavily grazed times hairy around the collar. Leaf sheaths are smooth ranges. Excessive use impairs its vigor, and over- to silky haired. Blades are flat to involute, character- grazed plants will die out during drought periods. istically curled or rolled inward with maturity, rough- Tillering is dependent on plant vigor during the previ- ened above and smooth below, 0.04 to 0.08 inch (1 to 2 ous year. Repeated cropping of black grama to 2 inches mm) broad, and 2 to 6 inches (5 to 15 cm) long. Ligules (5 cm) may prevent lateral spread. Damage may be are sometimes membranous at the base and often especially severe in overgrazed areas that have dete- fringed with hairs that are about 0.01 to 0.03 inch (0.2 riorated as a result of wind erosion (Canfield 1939). to 0.7 mm) long. Inflorescences usually consist of two Regeneration by stolon production can occur during short, comblike pedicellate spikes that range from 0.5 favorable years on conservatively grazed ranges to 2.0 inches (1.2 to 5.3 cm) in length and become sickle (Nelson 1934). Stolon production is negligible on ranges shaped at maturity (fig. 18). Each spike bears about 30 that are excessively grazed year after year. In addi- to 80 fertile spikelets, terminating with a single termi- tion, drought-induced mortality and competition from nal sterile spikelet. Fertile spikelets produce one per- less desirable and more competitive species can lead to fect floret and one or two sterile terminal florets. a loss of black grama plants and a reduction in ground Glumes are one nerved. The first glume is 0.08 to 0.14 cover and forage. inch (1 to 3.5 mm) long, linear, and translucent. The Black grama is seldom used in range seedings be- second is subequal to the lemma, 0.16 to 0.24 inch (4 to cause it is a slow starter and it is best adapted to harsh 6 mm) long, purplish, and three awned. The terminal environments. Heizer and Hassell (1985) recommended awn arises between two teeth and is 0.02 to 0.06 inch that black grama be seeded only to stabilize soil on (0.5 to 1.5 mm) long. Lateral awns are 0.04 to 0.08 inch critically eroding areas. Black grama has good poten- (1 to 2 mm) long with short, stiff hairs along the nerves. tial for ornamental use. The callus, a swollen node at the base of the lemma, is densely hairy (Arnow 1987; Hitchcock 1950). Spikes Varieties and Ecotypes become golden brown and tend to curve backward at As a result of low seed quality, high seed costs, and maturity or when the plant is suffering from drought. the difficulty of establishment, black grama has not Seeds do not disseminate rapidly at maturity. been seeded to any great extent. Two varieties are Blue grama produces an efficient, widely spreading available; little research has been conducted to de- root system (Briske and Wilson 1977). Numerous velop additional varieties. roots develop from short rhizomes. Fine fibrous roots

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sandy loam or rocky soils. It is often associated with galleta, sand dropseed, western wheatgrass, bluebunch wheatgrass, needle-and-thread, and broom snakeweed. Blue grama exhibits considerable variability in its growth habit and general appearance throughout its north-south range. In the north it tends to form a sod under favorable moisture conditions. In the south and at lower elevations it tends to form patches and not a complete cover. It is also a taller, more vigorous bunchgrass (Griffiths 1912). Blue grama normally remains dormant in spring, greening quickly with summer precipitation. It is peculiar in that it flowers from July through September. Because of its root system, blue grama is highly drought resistant (Ward and others 1990; Wilson and Briske 1979; Wilson and others 1976); it is one of the most important soil-binding grasses where adapted in the arid West. It has the ability to become dormant during drought periods, but resumes growth when temperatures and moisture conditions become favorable. It is a rapidly growing species, reaching maturity in 60 to 70 days (USDA Forest Service 1937). It produces highly palatable forage in late summer and early fall when other forage is scarce. Blue grama is resistant to grazing, but it is most productive when lightly grazed in late summer. It is highly fire tolerant, but it is not shade tolerant. Blue grama is adapted to a variety of soil types, but it most abundant on well-drained, sandy and sandy Figure 18—Blue grama spikes curve backward and loam soils of uplands. It can also be found on alkaline are golden brown when mature (Hitchcock 1950). and coarse-textured soils. In Arizona it is common on infertile, acidic soils (Nicholson and Bonham 1977). It is a common and important species on rocky outcrops, including mesas, canyon slopes, deep sandy soil on plains, and some midelevation grasslands.

Plant Culture Blue grama reproduces from seed, rhizomes, and are mostly concentrated near the soil surface, but tillers. Good seed crops are produced during years some extend deeper. with favorable moisture conditions. Seed quality is Ecological Relationships and Distribution generally low, varying with year and location. Wind, water, and animals disperse the seed (Albertson and Blue grama is a widely distributed climax species Weaver 1944). Adequate surface soil moisture is re- that occurs throughout the Great Plains from Alberta quired for a 2- to 4-day period to support germination and Manitoba south to Mexico, and from southern and initial seminal root growth. Lacking lateral semi- California eastward from southern Nevada and Utah, nal roots, seedlings die when the surface soil dries out Arizona, New Mexico, and Texas (Great Plains Flora before the seminal root elongates to deeper soil Association 1986; Hitchcock 1950; Stubbendieck and (Wicklow and others 1984). Critical factors for suc- others 1986). Blue grama is a major component of the cessful establishment of blue grama seedlings are short grass prairie. It is also found in scattered sites in seminal root initiation, good soil moisture for at least the Eastern United States. In the West, it is the most 6 weeks after emergence, and initiation of adventi- widespread and important species of its genus. It tious roots (Bock and Bock 1986; Albertson and Weaver occurs in grass and sagebrush communities on dry 1944). plains and foothills, and sometimes in pinyon-juniper The entire spikelet, including the awned fertile and ponderosa pine woodlands and mountain meadows floret and the awned and densely bearded rudimen- of the Intermountain area. At midelevations it grows on tary floret(s), is planted so the fertile florets are not

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 357 Chapter 18 Grasses separated during cleaning. These units are light and Varieties and Ecotypes fluffy and cannot be seeded through a conventional grain drill. Specially designed seed drills facilitate ‘Hachita’ was collected from an elevation of 4,000 ft uniform flow from the seed box to the soil. Shallow (1,220 m) in the Hachita Mountains of southern New planting on firm seedbeds improves seeding success. Mexico in an area with an average annual precipita- As a result of its wide distribution, blue grama is tion of 10 inches (250 mm). Hachita exhibits greater highly ecotypic. It is important to use seed sources drought tolerance and somewhat better seed and for- adapted to the edaphic and climatic conditions of the age production than ‘Lovington’. It is well adapted to area being seeded. areas in New Mexico, southeastern Colorado, and the panhandle area of Texas and Oklahoma that receive at Uses and Management least 8 inches (200 mm) of annual precipitation. It has been seeded with some success in the four corners area All classes of livestock and many wildlife species of Utah, Colorado, Arizona, and New Mexico (Alderson graze blue grama (Dittberner and Olson 1983). High- and Sharp 1994). quality forage, drought tolerance, and a desirable Lovington is a good forage and seed producer and is growth habit make it one of the most important range generally well adapted in areas of eastern New Mexico, grasses in areas where it is adapted. Although more southeastern Colorado, and northwestern Texas that palatable than most co-occurring species, blue grama receive at least 12 inches (300 mm) of annual precipi- has increased on many overgrazed ranges (Bock and tation. It has not been seeded to any extent in the Bock 1986; Weaver and Albertson 1956). Its persis- Intermountain West. tence is attributed to its vigorous root system, rhizomatous habit, and its ability to maintain adequate root carbohydrate reserves following grazing (Buwai Bromus anomalus and Trlica 1977). Following grazing, some lower leaves Nodding Brome ______generally remain near the soil surface and provide photosynthetic surface for plant recovery. With con- Synonyms tinuous heavy grazing and trampling, particularly on Bromus frondosus fine soils, blue grama may become sodbound and Bromus ciliatus var. montanus decline in forage production. Blue grama is a rapidly growing species and usually matures in 60 to 70 days. It is tolerant of grazing, but Bromus ciliatus yields the greatest return when grazed lightly in Fringed Brome______summer during its period of rapid growth. It can also be utilized in fall and winter. Grazing has often elimi- Synonyms nated the upright forms, leaving the less productive, sodforming types. This is especially true in the South- Bromopsis ciliata western United States. Bromus richardsonii Blue grama can be nutritious throughout the year. It greens up early and grows rapidly following summer Description storms. Its fine leaves are low in fiber and high in Nodding brome is a tall, nonrhizomatous perennial protein when green, making it an exceptional forage. existing as small tufts with culms 24 to 30 inches (6 to Protein and carbohydrate content are greatest during 10 dm) tall. Culms are typically puberulent, at least at early bloom, decreasing toward maturity. About 50 the nodes. Leaf blades are flat, 0.12 to 0.20 inch (3 to percent of its nutritive value, however, is retained at 5 mm) broad, sometimes slightly incurved or involute, maturity, making it an excellent fall or winter forage and smooth or sometimes with silky hairs. Ligules are (National Academy of Sciences 1971). In the South- small, 0.02 to 0.04 inch (0.5 to 1 mm) long. and slightly west, productivity of the shorter growth form of this irregularly toothed. Panicles are 2.8 to 5.5 inches (7 to species is relatively low. Growth of the shorter variety 14 cm) long, loose or open, and usually nodding. begins in May and June, providing very little green Spikelets are 0.67 to 1 inch (17 to 25 mm) long and forage during spring and early summer. slightly compressed. They produce 5 to10 flowers. Because of its drought hardiness and ability to Glumes are mostly minutely pubescent, or sometimes protect soils, blue grama is used extensively for protec- hairless. The first glume is 0.20 to 0.28 inch (5 to 7 mm) tion of critical eroding areas. It is also used for water- long and three nerved, or rarely one nerved. The shed stabilization and mined-land revegetation, as second glume is 0.25 to 0.31 inch (6.5 to 8 mm) long and well as for low maintenance turf plantings. three to five nerved. Lemmas are 0.35 to 0.47 inch

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(9 to 12 mm) long, five to seven nerved, rounded on the Bartos and Mueggler 1981; Bradley and others 1992a). back, and hairy. The awns are relatively short, about Nodding brome is more common in drier, more open 0.06 to 0.12 inch (1.5 to 3 mm) in length. communities in the mountain brush, pinyon-juniper, Nodding brome closely resembles fringed brome, ponderosa pine, and aspen types than fringed brome. and is known in some cases to hybridize with this Mueggler and Campbell (1986) reported that fringed species. Fringed brome is also a nonrhizomatous, brome is more common in shaded aspen and conifer tufted, native perennial with drooping or ascending communities, and nodding brome is more prevalent in branches (fig. 19). Arnow (1987) reported that al- open communities at lower elevations. Both species though both species occupy similar habits in Utah, occur as early, mid, and late seral species. They often they appear to be maintaining their integrity as dis- occur as dominant understory species in subalpine fir, tinct species. Only 3 percent of all specimens exam- Engelmann spruce, aspen, and blue spruce types ined exhibited any intermediacy. Nodding brome typi- (Alexander and Ronco 1987; Bradley and others 1992a; cally has pubescent glumes and awns are 0.04 to 0.16 Moir and Ludwig 1979). Both nodding brome and inch (1 to 4 mm) long. In comparison, the glumes and fringed brome are important species in aspen and culm nodes of fringed brome are glabrous, and its awns conifer forest types including open meadows and grass- range in length from 0.08 to 0.24 inch (2 to 6 mm) long lands. In addition, nodding brome is equally impor- (Arnow 1987). tant in mountain brush, pinyon-juniper, and sagebrush types, occurring in various shrub and herb Ecological Relationships and Distribution associations (Allman 1953; Bartos and Mueggler 1981). Fringed brome is commonly encountered on moist to Both nodding brome and fringed brome range from seasonally dry habitats, including shaded and open British Columbia and Saskatchewan south through the parks. It grows well on semiwet or moist soils in valley Western United States and Mexico, and east to the bottoms and meadows. It is adapted to poorly drained Dakotas and Texas (Arnow 1987; Cronquist and others conditions (Mattson 1984), but also grows on well- 1977; Hitchcock 1950; Hitchcock and others 1969). In drained and rocky substrates (Butterwick and others the Intermountain region, nodding brome and fringed 1992; Crouch 1985). brome are found on overlapping sites. Both occur on a variety of habitat types including dry to moderately Plant Culture moist sagebrush, mountain brush, aspen, spruce-fir, ponderosa pine, lodgepole pine, mountain herblands, Nodding brome and fringed brome have not been grassy meadows, and riparian communities at 5,460 to extensively used in restoration programs. Limited test- 10,890 ft (1,680 to 3,350 m) elevation (Arnow 1987; ing has been conducted to evaluate either species for range or watershed plantings. Both species exhibit similar characteristics in relation to seed germination and stand establishment. Seed viability and germination of wildland collections are normally quite high. Seeds of both species can be harvested, cleaned, and seeded as easily and successfully as other species of brome. Plummer and others (1968) reported that seeds of nodding brome germinate easily without special treatment. Hoffman (1985) found that seeds of fringed brome are nondormant, and nearly all seeds germinated with conventional treatments. Initial establishment and growth of both species is normally greater than for smooth brome. Plants of both species tend to form small clumps, and do not persist as long as sodforming species of brome. Both nodding brome and fringed brome usually form full stands when seeded alone, even on irregular terrain. They provide more open stands than sodforming bromes, including smooth brome. Nodding brome plantings established in central Utah persisted as nearly solid stands for 10 to 30 years, but were slowly invaded by associated grasses and broadleaf herbs. Figure 19—Nodding brome forms small, Although stands decreased in density, the grass re- nonrhizomatous tufts with loose, open, panicles mained important and spread by natural recruitment (Hitchcock 1950).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 359 Chapter 18 Grasses to adjacent openings. Natural spread of both nodding and communities. Where the distributions of these two fringed brome is quite successful, equaling that of most bromes overlap, mixed plantings are recommended. Al- perennial grasses adapted to mountain communities. though neither nodding brome nor fringed brome form Seeds of both nodding brome and fringed brome are dense sod, both establish quickly and provide satisfac- large and easily seeded through most drills. Seedlings tory cover. Both prefer deep and rich soils, but they are of both species also establish well from broadcast also encountered on shallow, well-drained soils. distribution and harrowing to cover the seed. Seeds Nodding brome and fringed brome have been grown can be aerial seeded under aspen, oak, or other shrubs to a limited extent in hay meadows, providing yields as and trees prior to leaf fall. Planted in mixtures or high as 2 tons per acre (2,250 kg per ha) (Dayton 1931). alone, both species can be seeded at rates used for Seed production fields have not been established for smooth brome, between 2 to 4 pounds per acre (2.2 to either species, thus large seedings are not feasible. 4.5 kg per ha). Observation and management of wildland stands sug- Nodding brome and fringed brome are highly com- gest both species could be grown under cultivation patible with other species and can be seeded in mix- without special care. Both species grow well in associa- tures without suppressing establishment or growth of tion with other herbs, and are useful grasses for associated herbs. Both species are less competitive restoring disturbed watersheds. Plants establish than smooth brome and could be planted in areas quickly from direct seeding, and spread to stabilize where smooth brome has been recommended. small openings. Neither species is invasive; both can be seeded in mixtures with native broadleaf herbs to Uses and Management restore complex communities. Both nodding brome and fringed brome should be Both species furnish excellent forage and ground utilized in restoration plantings, and could be used to cover. They are rated as among the best forage grasses replace smooth brome where reestablishment of na- on Western rangelands (Dayton 1931). Plants are tive communities is required. To date, ecotypes of grazed readily in spring and summer by livestock, these species have not been identified, but seed should deer, and elk (Humphery 1960). The foliage of both be collected and planted within areas of occurrence. species can become tall and rank at maturity and is Moving high-elevation collections to lower and drier less palatable in late summer. However, both species sites is not advised. can be removed by close grazing (Humphrey 1960). Livestock and wildlife graze seedheads before and Varieties and Ecotypes after the seeds have matured. New basal leaves are produced in fall and selectively grazed. Domestic sheep There are no releases. seek the plants in mid and late summer as seeds mature and after fall greenup. Nodding brome and fringed brome have often been Bromus carinatus misidentified and planted in areas where they are not Mountain brome, California adapted. Nodding brome is better suited to open aspen brome ______communities and sites at lower elevations. It can be planted in the upper sagebrush and mountain brush Synonyms types to provide forage for wildlife. This species should be more widely used to plant oak brush sites following Ceratochloa carinata wildfires to control erosion and reestablish a native Bromus marginatus understory. It can also be planted in pinyon and Bromus breviaristatus juniper restoration projects where it is native. Nod- Bromus polyanthus ding brome is not as productive as smooth brome or mountain brome. It is less palatable than smooth Description brome, but equal to mountain brome. Fringed brome Mountain brome is a member of a highly polymor- normally grows at somewhat higher elevations than phic complex of related taxa (Arnow 1987). It has nodding brome, and it is better adapted to shaded sometimes been recognized as three or more separate communities. It is also more common in tall forb types species (Hitchcock 1935), usually including B. at high elevations. marginatus, B. polyanthus (both perennials with awns Nodding brome and fringed brome can be planted on less than 7 mm long), and B. carinatus (an annual or open disturbances, including forested sites and moun- biennial with awns more than 7 mm long). B. tain brush fields burned by fires. They can be mixed with marginatus is pubescent throughout, while the spike- other species, including natives, and planted by lets of B. polyanthus are glabrous (Cronquist and aerial seeding to control erosion and restore native others 1977). B. marginatus and B. carinatus are

360 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses morphologically quite variable but scarcely distinct Ecological Relationships and Distribution from each other. B. marginatus interbreeds with both B. carinatus and B. polyanthus (Wasser 1982). Plants Mountain brome is widespread in North America, with characteristics of all three species can sometimes occurring from Alaska and Canada south to Califor- be found within a single population. Hitchcock and nia, Texas, and most of Mexico. It has been introduced others (1969) concluded there was no basis for the in some Midwestern States and naturalized in Europe division of these taxa based on their annual or peren- (Arnow 1987; Cronquist and others 1977; Hitchcock nial growth habits. The following description of B. 1950; Hitchcock and others 1969). Mountain brome is carinatus follows Arnow (1987) and Cronquist and common in both early and late seral communities. It others (1977) and includes these taxa. occurs in the subalpine fir-Engelmann spruce zone Mountain brome is a tall, short-lived native peren- and in high-elevation sedge meadows (Tisdale and nial that often flowers during its first growing season Bramble-Brodahl 1983). In the Intermountain area, it (fig. 20). Culms are solitary or tufted, usually erect, 1.0 is commonly found in aspen and conifer communities to 2.3 ft (6 to 14 dm) tall, and smooth to minutely hairy. and in associated herblands and meadows (Ellison Leaf blades are flat, 0.08 to 0.55 inch (2 to 14 mm) wide, 1954; Mueggler and Campbell 1986). It also occurs in rarely involute, and smooth to roughened or silky mountain big sagebrush communities. It is more com- haired. Ligules are 0.04 to 0.12 inch (1 to 3 mm) long, mon on cooler sites (Mooney 1985) in the lower eleva- smooth or with silky hairs, and irregularly serrate tions of these communities (Hironaka and others 1983; along the margins. Panicles are usually 4 to 12 inches Tisdale and Bramble-Brodahl 1983). (1 to 3 dm) long, loosely contracted, and nodding. Spikelets vary from one to four per branch. They are Plant Culture 0.6 to 2.4 inches (1.5 to 6 cm) long, strongly com- Mountain brome reproduces primarily by seed pressed, and produce 4 to 16 flowers. Glumes are (USDA Forest Service 1937), although some tillering lanceolate and strongly keeled. The first glume is 0.27 to occurs (Stubbendieck and others 1992). Plants flower 0.43 inch (7 to 11 mm) long and three- to five-nerved. from late May to early August. Harlan (1945) reported The second glume is 0.35 to 0.51 inch (9 to 13 mm) long that cross-fertilization is most common when growing and five to seven nerved. Lemmas are 0.43 to 0.67 inch conditions are optimum, while adverse conditions (11 to 17 mm) long, carinate keeled, and smooth to stimulate self-fertilization. In California, both open roughened. The apex is slightly notched to entire. pollinating and cleistogamous florets are commonly Awns are 0.16 to 0.31 inch (4 to 8 mm) long. Anthers found on the same plant (Hickman 1993). are small and included within the lemma. Native stands frequently produce heavy seed crops, and a limited amount of wildland-harvested seed is sold each year. Agricultural production is required to meet the demands for seed of this species, and highly productive mountain brome seed fields have been established on low-elevation farmlands. Seeds are harvested by combining, and yields average about 250 pounds per acre (280 kg per ha) (Smith and Smith 1997). There are about 103,700 seeds per pound (228,000 per kg) (Smith and Smith 1997). Seed germination of mountain brome is relatively high, normally exceeding 85 percent. Plummer (1943) reported a higher percentage of seeds germinated in a shorter time when exposed to alternating rather than constant temperatures during germination. Paulsen (1970) found that optimum germination occurred over a wide range of temperatures. Satisfactory wildland stands can be established from either fall or spring plantings in weed-free areas. When planted alone, a seeding rate of 4 to 10 pounds per acre (4.4 to 11.2 kg per ha) is recommended. Seeds Figure 20—Common on many high-elevation sites, should be planted at depths of 0.25 to 0.50 inch (0.6 to mountain brome occurs in subalpine, mountain 1.3 cm) (Jensen and others 2001). Seeds are large and meadow, aspen, conifer, and mountain big sage- can be planted with most drills or broadcast seeders. brush communities (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 361 Chapter 18 Grasses

Broadcast seed should be covered by light harrowing. others 1992). Seedheads are used by birds and small This species establishes better than many other grasses mammals. The herbage becomes coarse and fibrous at when broadcast seeded. Mountain brome is frequently maturity. Regrowth in summer is moderate, providing seeded in mixtures with other native species. It is also late fall forage. Fall regrowth is fair. Mountain brome seeded with introduced species to provide ground is tolerant of grazing, but excessive use weakens the cover and quickly stabilize severely disturbed sites. stands (Stubbendieck and others 1992). Consequently, the seeding rate for mountain brome Mountain brome is tolerant of wildfires (Brown and may be adjusted so that it will provide adequate DeByle 1989), recovering from seed and surviving ground cover, but also permit establishment of other plants. Stands may be reduced initially, but they seeded species (Sours 1983). Seedings should be pro- recover to prefire levels after a few years. Success of tected from grazing until plants are well established postfire seedings of this species has been variable. and seedheads are produced. Established seedings are useful for stabilizing soil and providing cover and forage, but they may prevent Uses and Management recovery of native species or coseeded natives (Steele and Geier-Hayes 1993). Seedlings of mountain brome are vigorous and establish quickly, but plants may not produce seed for 2 Varieties and Ecotypes to 3 years (Cronquist and others 1977). Stands of mountain brome generally consist of scattered plants, ‘Bromar’ mountain brome was developed from na- but they sometimes develop a dense sod. Mountain tive material collected near Pullman, WA (Alderson brome can increase in density when growing with and Sharp 1994). It is adapted to streambanks, semiwet other perennials. Plants, however, are short lived and meadows, subalpine conifer forests, aspen, and upper must be allowed to set seed every 2 or 3 years to permit mountain brush communities in the Intermountain natural reseeding. If natural seeding does not occur, West. Bromar has been widely used to stabilize log- the species can be replaced by longer lived species. ging, mining, roadway, and other disturbances. It does Hafenrichter and others (1949) reported that longev- not spread aggressively, and it is compatible with ity and persistence are important characteristics to other seeded or indigenous species. Livestock and big consider when selecting plant material of this species. game use its forage, especially prior to the production Mountain brome is seeded as an early successional of seedstalks. This variety exhibits good seedling vigor species and establishes quickly on disturbed and weedy and spring greenup. It is, however, susceptible to head sites at mid to upper elevations. It is seeded alone or in smut and fairly short lived (6 to 10 years). Bromar is mixtures on depleted livestock and game ranges, burns, a good seed producer; stands reseed naturally if not roadways, and mined lands in forested areas. The grazed excessively. Bromar should be fall seeded by species is effective for providing rapid, cold-tolerant drilling or broadcasting cover and erosion control in moderate to high eleva- ‘Garnet’ mountain brome germplasm, released in tion watersheds, as it grows rapidly and produces 2000 by the Upper Colorado Environmental Plant deep, well-branched root systems (Stevens and others Center, Meeker, CO (Englert and others 2002), origi- 1992). Mountain brome is also an excellent grass for nated from material collected near Garnet in Powell pasture or hay (Sours 1983). County, MT (Alderson and Sharp 1994), at an eleva- Mountain brome tolerates a wide variety of soil tion of 5,800 ft (1,770 m). Garnet has been evaluated textures, but it most often occurs on medium-textured throughout the Intermountain West. It exhibits good to loamy soils in the pH 5.5 to 8.0 range (Sours 1983). stand longevity and excellent head smut resistance, It is most productive on deep, rich, moist soils, but it two characteristics lacking in many populations of can grow on dry, shallow, and infertile soils on dis- this species. Garnet is recommended for seeding on turbed sites. The species is winter hardy, moderately deep, fertile, mesic, medium-to fine-textured soils in tolerant of shade and salinity, and weakly to moder- areas receiving at least 15 inches (380 mm) of annual ately tolerant of drought and heat (Hassell and others precipitation. It thrives in climates with cool, dry 1983; Steele and Geier-Hayes 1993). It does not en- summers and good winter precipitation. It does not dure long periods of flooding or high water tables. tolerate high water tables or flooding, but it does Mountain brome is palatable to livestock and big exhibit good shade tolerance and fair fire tolerance, game (Stubbendieck and others 1992). It produces and it is very winter hardy. Garnet is adapted to moderate amounts of high-quality herbage and re- mountain brush and mountain big sagebrush commu- ceives considerable use in spring and early summer nities. It is also seeded in openings in aspen and prior to seed dispersal (Hassell and others 1983). Elk conifer forests and in subalpine areas. It is not aggres- use it in summer (Leege 1984; Stubbendieck and sive and can be seeded in mixtures.

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Bromus inermis it has been planted on overgrazed rangelands, roadways, ski areas, mined lands, burned areas, unstable Smooth Brome, Hungarian watersheds, and other disturbances at mid to upper Brome ______elevations (fig. 21). It is now distributed from Canada to Southern California and east to New Mexico and Synonyms much of the North-Central and Northeastern United Festuca inermis States (Arnow 1987, 1993; Cronquist and others 1977; Bromopsis inermis Hitchcock 1950; Hitchcock and others 1969). Bromus pergans var. purparescens Smooth brome is adapted to streambanks, valley Bromus pergans var. longispicata bottoms, and upland sites (Hardy BBT Limited Bromus pumpellianus 1989) in exposed to partially shaded areas that Bromus ciliatus var. coloradensis receive 14 to 20 inches (356 to 508 mm) of annual precipitation (Jensen and others 2001; Plummer Description and others 1968) and sites that are subirrigated or intermittently wet. It grows on a variety of well- Smooth brome is a long-lived, strongly rhizomatous, drained, medium-textured clay loam to sandy loam sodforming perennial with the few culms about 5.9 to 17.7 inches (1.5 to 4.5 dm) tall. Leaf sheaths are smooth to silky haired. Blades are flat and broad, about 0.16 to 0.51 inch (4 to 13 mm) wide, rarely involute, and hairless or rarely silky haired on the upper side. Leaf margins and nerves are rough and hairy. Ligules are small, about 0.06 to 0.20 inch (1.5 to 5.0 mm) long, and irregularly serrate or reduced to a marginal fringe of minute hairs. Panicles are 2.8 to 9.1 inches (7 to 23 cm) long, moderately open, oblong, and erect. Panicle branches are ascending; the upper ones are appressed. Spikelets are 0.6 to 1.3 inches (1.6 to 3.2 cm) long, 4 to 10 flowered, very narrow and round when young, but becoming somewhat compressed with age. Glumes are hairless and smooth. The first glume is 0.18 to 0.31 inch (4.5 to 8.0 mm) long, narrowly lance shaped, and one to three nerved. The second glume is 0.24 to 0.39 inch (6 to 10 mm) long, lance shaped, and three to five nerved. Lemmas are 0.35 to 0.47 inches (9 to 12 mm) long, rounded on the back, smooth or rarely minutely hairy, and greenish to purplish tinged. The lemma apex is bidentate to slightly rounded. Awns, when present, are up to 0.08 inch (2 mm) long. Smooth brome flowers from June to August, and sometimes into early September. Smooth brome hybridizes readily with native B. pumpellianus, a taxon that Elliott (1949) suggested might be reduced to a subspecies of B. inermis due to extensive introgression. Arnow (1987) stated that she had never seen material that could be identified as belonging to B. pumpellianus, which is distinguished from B. inermis by its pubescent culm nodes and lemmas.

Ecological Relationships and Distribution Figure 21—A strongly rhizomatous introduced Smooth brome is a European introduction that has species, smooth brome receives wide use for wa- been widely seeded for hay and pasture in irrigated tershed stabilization and postfire seedings on for- and nonirrigated fields. In the Western United States ested sites, but it does preclude recovery of native species (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 363 Chapter 18 Grasses soils of at least moderate depth; it is less productive Grasshoppers sometimes damage or destroy develop- on fine or coarse soils and on shallow soils. Smooth ing stands. Mature plants spread vegetatively, while brome is cold tolerant; tolerant of poorly drained spread from seed is moderate (Plummer 1977). Al- soils, saline or acidic conditions, and drought; but it though stands may require 3 years to become fully does not tolerate high alkalinity (Coulman 1987; established, they are aggressive and may dominate Hafenrichter and others 1968; Wambolt 1976). seedings or recovering native species. Because of its Smooth brome becomes dormant in winter, and ability to spread vegetatively, smooth brome plants initiates growth in early spring (Smoliak and others are long lived. Plantings have persisted for 60 years or 1981). Plants are productive, developing abundant more (Plummer and others 1968). Established stands vegetative growth and large seedheads (Jensen and of the species invade and dominate local disturbances others 2001). They become dormant during dry peri- (Boggs 1984; Boggs and Weaver 1992; D’Antonio and ods, but initiate regrowth with adequate precipitation Vitousek 1992). in fall. Following wildfires, smooth brome regenerates by Uses and Management sprouting from rhizomes and tilllering. Productivity is increased by early- or late-season fires that remove Smooth brome is one of the most commonly seeded litter, particularly in sodbound stands (Higgens and exotic grasses for pastures and wildlands in North others 1989; Hughes 1985). Fires occurring in late America (Mitchell 1982, 1987b). Selected varieties can spring are generally more damaging (Masters and be used on a wide range of irrigated and nonirrigated Vogel 1989). sites with varying soil conditions. A number of smooth brome varieties with differing Plant Culture areas of adaptation have been developed to provide erosion control on disturbed lands and nutritional Smooth brome is generally a cross-pollinating spe- forage or hay when planted alone or in combination cies (Knobloch 1944), but self-fertilization can occur with legumes. Smooth brome regrowth following cut- between flowers in different spikelets (McKone 1985). ting is slow. Sixteen to 18 inches (41 to 46 cm) of annual Because of insect herbivory, seed set is lower for precipitation are generally required to maximize for- smooth brome than for many other brome species age production. Pastures and fields that become (McKone 1985; Nielson and Burks 1958). Seeds ma- sodbound require application of fertilizer and cultiva- ture in early to midsummer. They are harvested by tion for renovation (Jensen and others 2001). combining and cleaned with an air screen machine. Smooth brome has been planted in a wide variety of There are 136,100 to 149,700 seeds per pound (300,000 plant communities to provide palatable forage for to 330,000 per kg) (AOSA 1999). Smooth brome seed wildlife and livestock. Seeded stands, particularly has remained viable for periods of 22 months to more those on fertile soils (Currie and Smith 1970), are than14 years in storage (Hafenrichter and others palatable and tolerant of heavy use. The species has 1968; McAlister 1943; Rincker and Maguire 1979). been seeded in saltgrass, pinyon-juniper, quaking The AOSA (1999) rule for testing germination of smooth aspen, and subalpine communities (McGinnies brome stipulates an incubation temperature alternat- 1960a,b, 1975; Sampson and others 1951). ing between 68 and 86 ∞F (20 to 30 ∞C) (16 hrs/8 hrs). Wildlife use of smooth brome varies among animals A moist prechill at 41 to 50 ∞F (5 or 10 ∞C) for 5 days is and season. Deer browse the species (Stubbendieck recommended for fresh or dormant seed. and others 1992); elk take it primarily in winter Smooth brome should be planted in late fall or early (Hobbs and others 1981). Ducks and upland birds use spring at depths of 0.25 to 0.75 inch (0.6 to 1.9 cm), it for nesting cover, while geese graze the new shoots. depending on soil texture, but seedlings do emerge Rodents eat the forage and seeds. from greater depths (Plummer 1943). Seeds may be The rhizomatous habit and aggressive growth of drilled or broadcast and covered. The elongate seeds smooth brome make the species a valued erosion sometimes plug drills (Jensen and others 2001). Seed- control plant (Hardy BBT Limited 1989). It is used for lings can germinate at low temperatures under snow stabilization of degraded streambanks, mined lands, (Bleak 1959) and develop rapidly as temperatures logged areas, and other disturbances (Elliott and oth- increase in spring. Smooth brome requires fertile soil. ers 1987; Ward and others 1986). Loss of native veg- Fertilization or seeding with a nitrogen fixer may be etation, however, may occur because of the aggressive- required to maintain stands when natural soil fertility ness of this species (Smoliak and others 1981). is inadequate (Jensen and others 2001). Seedlings are Smooth brome has been widely used to hold surface not strong, but once established they grow rapidly soil and produce forage rapidly following wildfires (Hardy BBT Limited 1989; Hassell and others 1983). (Clary 1988; Crane and Habeck 1982). Because

364 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses recovering native grasses, forbs, and shrubs are often Bromus riparius as effective in holding the soil and controlling erosion and because the aggressiveness of smooth brome may Meadow Brome ______preclude recovery of native species, its value for Synonyms postfire seedings has come under question (Clark and McLean 1979; Crane and Habeck 1982). Bromus erectus Bromus biebersteinii Varieties and Ecotypes Bromus macounii

Smooth brome varieties are generally divided into Description three types: southern, intermediate, and northern. Southern varieties include ‘Lincoln’, ‘Saratoga’, Meadow brome, earlier described as Bromus erectus ‘Baylor’, ‘Beacon’, ‘Tempo’, ‘Signal’, ‘York’, ‘Radisson’, (Hitchcock 1950), is now recognized as Bromus riparius ‘Badger’, ‘Achenbach’, and ‘Southland’. Intermediate (Vogel and others 1996). Based on descriptions pro- forms include Tempo, ‘Magna’, and Signal. ‘Manchar’, vided by Tsvelev (1984), Vogel and others (1996) con- ‘Carlton’, ‘Elsberry’, ‘Polar’, ‘Jubilee’, ‘Bravo’, and ‘Re- cluded that most North American introductions and bound’ are northern forms (Alderson and Sharp 1994). cultivars of B. biebersteinii are also B. riparius. Meadow Southern varieties are strongly rhizomatous brome is an introduced, long-lived, cool-season, peren- spreaders that are not compatible with natives. nial bunchgrass. Culms are tufted, 2 to 3 ft (6 to 9 dm) Their long, wide, and somewhat coarse leaves are tall, erect, slender, and glabrous to silky haired. Leaf primarily basal, but some occur along the culm. The blades are narrow and sparsely hairy. Ligules are 0.06 panicles are somewhat contracted with green to inch (1.5 mm) long. Panicles are narrow and 4.0 to 7.8 light-red spikelets. Southern varieties generally inches (10 to 20 cm) long with ascending to erect exhibit good seedling vigor. They require less mois- branches. Spikelets are usually 5 to 10 flowered. ture than do the northern types, and they can be Glumes are tapered to a point, the first is about 0.24 to planted in areas receiving full sunlight or consider- 0.31 inch (6 to 8 mm) long, and the second about 0.31 able shade. They green up early in spring and are to 0.39 inch (8 to 10 mm) long. Lemmas are 0.39 to 0.47 good forage producers. inch (10 to 12 mm) long and smooth or evenly rough Northern varieties are also aggressive spreaders hairy on the back. Awn length is 0.20 to 0.24 inch (5 to that are not compatible with natives. Their long leaves 6 mm) (Hitchcock 1950). are intermediate in width, soft, smooth, more or less Meadow brome is somewhat similar in appearance vertical, and occur well up the culms. Panicles are to smooth brome (fig. 22). The main difference is the open with red to purple spikelets. Northern varieties exhibit good seedling vigor. They require more moisture than southern varieties, begin growth about 5 to 10 days later, and are generally less productive. Lincoln and Manchar are two of the most commonly seeded smooth brome varieties in the Intermountain West. Lincoln, a southern variety, is derived from material thought to be of Hungarian origin (Alderson and Sharp 1994). It is seeded in aspen, mountain brush, ponderosa pine, upper pinyon-juniper, and big sagebrush communities that receive 13 or more inches (330 mm) of annual precipitation. It is also well adapted to semiwet meadows and streambanks. Lincoln establishes quickly from drilling or broadcasting and is a very aggressive spreader that eventually dominates mixed seedings. Manchar, a northern variety from Manchuria, China (Alderson and Sharp 1994), is adapted to openings in aspen stands, conifer forests, mountain big sagebrush communities, semiwet meadows, and riparian areas. It is less well adapted to mountain brush communities. Manchar is aggressive and will dominate seeded Figure 22—Meadow brome can be distinguished areas, especially those that are heavily grazed. from smooth brome by the pubescence on the awns and leaves. Its panicles rise well above the leaves (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 365 Chapter 18 Grasses pubescence on the awns and leaves of meadow brome Instead, both smooth brome and meadow brome have (Knowles and others 1993). In addition, its rhizomes been used to provide effective ground cover and fur- are much shorter than those of smooth brome, and its nish palatable forage. Use of meadow brome in the stems and seeds are more pubescent (Vogel and others West began in 1966 with the release of ‘Regar’ for 1996). planting as a forage and conservation species on rangelands and to provide watershed stabilization (Casler Ecological Relationships and Distribution and Carlson 1995; Knowles and others 1993). Normally long lived, meadow brome is less winter Meadow brome is a European introduction hardy and drought tolerant than smooth brome (Weintraub 1953), and has been planted or has estab- (Knowles and others 1993). It has been widely and lished in localities from Maine to New York, as well as successfully planted in irrigated pastures for grazing in the Pacific Northwest and the South (Hitchcock and seed production. It is also well-suited to mountain 1950). In recent years it has been seeded on range and brush, aspen, conifer forests, and mountain park com- watershed sites in the West. Meadow brome has proven munities, and it is recommended for planting in inter- adapted to mountain brush, ponderosa pine, aspen, mediate and high-elevation woodlands (Hansen 1972). and subalpine communities in the Intermountain re- In western Montana it has competed more success- gion. It is well-suited to acidic soils of the Idaho fully with cheatgrass than has crested wheatgrass. It Batholith, including disturbances where subsoils have is less well adapted to more arid regions occupied by been exposed. This species is marginally adapted to big sagebrush and associated shrubs and bunchgrasses. drier regions occupied by pinyon-juniper and big sage- Meadow brome is not as productive as smooth brome, brush communities. In the Intermountain region, the but it is still regarded as an important forage species. plant has not persisted on sites receiving less than 12 Spring greenup is rapid (Schwendiman and others to 14 inches (30 to 36 cm) of annual precipitation. 1946), and plants provide excellent early spring and summer herbage for game and livestock. Regrowth is Plant Culture more rapid for meadow brome than for smooth brome (Knowles and others 1993); consequently, pasture and Seeds of meadow brome are similar in size to those rangeland stands supporting the species recover well of smooth brome and other brome species. Knowles from early-season grazing. When seeded on wet and and others (1993) reported that compared to other semiwet sites in mountain communities this grass bromes, the seeds of meadow brome shatter more also provides excellent summer and fall herbage, and easily and require more processing to remove the awns plants persist well. Herbage quality diminishes as and pubescence to assure that seeds can be seeded plants mature, but the foliage remains quite palatable uniformly. Meadow brome establishes quickly, and even though the leaves are quite pubescent (Hansen dense, uniform stands generally develop. Plants grown 1972). This species provides a valuable hay crop, even under cultivation produce reliable and abundant seed on soils with low fertility (Hansen 1972). Rangeland crops in most years. Successful stands have estab- plantings have not diminished in productivity or be- lished on range and wildland sites in the West by drill come sodbound due to a decrease in available nitrogen, seeding or broadcasting and harrowing to cover the as is commonly experienced with smooth brome. seed. When drill seeded alone, planting rates of 8 to 10 Vegetative spread is not so aggressive for meadow pounds per acre (8.9 to 11.2 kg per ha) are sufficient. brome as for smooth brome. Plantings on high-eleva- The rate should be decreased to 2 to 4 pounds per acre tion mountain herblands and in mountain brush com- (2.2 to 4.4 kg per ha) when seeded in mixtures with munities have spread little from the original rows over other species. Rates should be increased slightly for a 25-year period. In addition, established stands have seedings on severely disturbed and eroding sites to not seriously reduced the presence of associated na- assure rapid development. Plants provide good ground tive herbs. This grass is much less competitive with cover in the first and second seasons. Like mountain native species than smooth brome, and could be planted brome, meadow brome is susceptible to head smut, in areas where it is important to maintain the compo- and seeds should be treated prior to planting to reduce sition of native plants. the chance of infestation.

Uses and Management Varieties and Ecotypes Regar is an early maturing bunchgrass that is Meadow brome and smooth brome are the two spe- adapted to areas in upper pinyon-juniper, mountain cies of brome cultivated for grazing and seed produc- brush, and aspen communities that receive 14 inches tion in pastures in North America (Vogel and others (356 mm) or more of annual precipitation. It was 1996). Although there are native bromes in the West, developed from material collected at Zek, Turkey few have been developed for restoration plantings.

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(Alderson and Sharp 1994), and released by the Aber- rounded on the back. Their margins are straight, and deen Plant Materials Center in Aberdeen, ID (Foster they are acute to awn tipped apically. The keel is and others 1966). This variety is characterized by good smooth to variously pubescent. The first glume is 0.20 drought tolerance, excellent winter hardiness, and to 0.25 inch (5 to 6.5 mm) long, keeled, and acute or moderate spread. Plants greenup early in spring. tapered to an awnlike tip that is about 0.08 inch (2 They grow well in full sunlight and partial shade. They mm) long. The second glume is 0.12 to 0.24 inch (3 to are leafy and exhibit good regrowth, particularly in 6 mm) long and membranous. Florets disarticulate midsummer. Cooper and others (1978) found that above the glumes, which remain attached to the spike- yields were greater for Regar than for smooth brome let. Flowering occurs from June through August (Arnow when seeded in grass-legume mixtures. Regar may be 1987, 1993; Cronquist and others 1977; Hitchcock planted in fall or early spring when seeds germinate 1950; and Hitchcock and others 1969). and establish readily. It does not spread as aggressively as smooth brome, and it is somewhat compatible Ecological Relationships and Distribution with other recovering natives. It is seeded in the Intermountain West for forage and for stabilization of A European introduction, orchardgrass is widely disturbed sites. distributed throughout North America. In the West it Subsequent meadow brome cultivar releases, ‘Fleet’ is found in sagebrush, mountain brush, ponderosa and ‘Paddock’, were developed by Agriculture Canada. pine, aspen-fir, and occasionally in desert shrub com- Seed production of both cultivars generally exceeds munities. Orchardgrass was introduced as a hay and that of Regar by more than 65 percent (Vogel and pasture grass, and in the West, it is commonly grown others 1996). in irrigated areas or on sites receiving a minimum of 18 inches (457 mm) of annual rainfall. Although it generally occurs as a seeded species, orchardgrass Dactylis glomerata volunteers on moist nonirrigated sites (Vallentine Orchardgrass______1985). It requires well-drained, medium-textured, deep to moderately deep soils that are moderately acid to Synonyms moderately alkaline. It will grow on shallow, gravelly, or stony soils (Vallentine 1961). Orchardgrass ini- Bromus glomeratus tiates growth in early spring, and it is highly produc- Festuca glomerata tive during the cool season. Due to its deep rooting habit it is also capable of considerable summer growth Description under favorable conditions (USDA Natural Resources Attempts have been made to place orchardgrass in Conservation Service 2000). Orchardgrass produces different generic complexes, but for the most part, palatable, nutritious forage, and it tolerates heavy taxonomists in the United States and Europe agree grazing. It is reported to increase or remain stable that it should remain in the monotypic genus Dactylis. after burning (Cocking and others 1979). The species is a tufted perennial that grows 1 to 4 ft (3 to 12 dm) tall and produces short rhizomes. Leaf Plant Culture sheaths are shorter than the internodes. Ligules are Orchardgrass seed matures evenly and is ready for 0.16 to 0.31 inch (4 to 8 mm) long and oblong to ovate harvest in mid-August (USDA Natural Resources with irregularly lacerated margins. Leaf blades are Conservation Service 2000). There are 480,000 to 0.16 to 0.39 inch (4 to 10 mm) wide, lax or loosely 650,000 seeds per pound (1,058,200 to 1,432,990 per arranged, and roughened on the surface. Panicles are kg). Orchardgrass can be spring seeded, but later 1.8 to 7.9 inches (4.5 to 20 cm) long, laterally com- dates can be successful with adequate irrigation. Fall pressed, and mostly interrupted. The primary panicle seedings may be more appropriate on sites that are branches, one to three at each node of the rachis or axis likely to dry early in the season. A clean, firm, weed- of the panicle, measure about 1.2 to 4.7 inches (3 to 12 free seedbed is recommended. Seed should be planted cm) long or longer, and are erect to widely spreading or about 0.5 inch (1.3 cm) deep. Seed increase fields reflexed. The branches are naked below, but with one should be planted with the rows at 28- to 40-inch to several dense clusters of subsessile to short- (71- to 102-cm) spacings. Orchardgrass establishes pedicillate spikelets borne near the branch tips and well in mixtures with other competitive introductions arranged on one side of the axis. Spikelets are 0.20 to where adequate precipitation, at least 12 inches 0.35 inch (5 to 9 mm) long, two to five flowered, and (305 mm), is received annually (Hardy BBT Limited occasionally purple tinged. Glumes are subopposite, 1989). lance shaped, often asymmetrical, and keeled to

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Uses and Management of orchardgrass should be used to avoid losses from winterkill (Vallentine 1961). Orchardgrass is used in seed mixes with other grasses Vallentine (1961) reports that all classes of livestock and forbs for revegetating overgrazed or otherwise readily use orchardgrass, and deer, elk, and bighorn disturbed lands (Hardison 1980) (fig. 23). Success of sheep graze it. The basal rosette of the plant is green these seedings depends on the selection of appropriate through fall and into the winter and provides a good cultivars. Orchardgrass has been planted in logged source of forage for wildlife (Dalke 1941). Wildlife use over or burned areas, and it provides cover and forage of seedings on burns and other disturbances increases for livestock and wildlife (Clary 1975a). It has also been when orchardgrass is seeded (Evanko 1953; Leege and used for mine reclamation, ski slope stabilization, and Godbolt 1985; Lowe and others 1978). Seedings also suppression of annual weeds (Behan 1983; Hardy 1989). provide habitat for rodents (Reynolds and Johnson Orchardgrass is highly shade tolerant and has been 1964) and nesting and brood rearing habitat for upland seeded under aspen and open conifer stands. birds as well as escape and winter cover. It is used by Orchardgrass is recommended for moderate early- rabbits and Canada geese (Crawford and others 1969). season grazing (Humphrey 1960). It provides excel- Although orchardgrass is widely seeded following lent, highly digestible forage (Vallentine 1961). Rota- wildfires or other disturbances, it interferes with tional grazing practices are best for irrigated pastures. natural regeneration of conifer and native understory Orchardgrass is grown with alfalfa for hay crops species and with establishment of planted conifers because of its ability to recover from grazing or clip- (Anderson and Brooks 1975; Elliott and White 1987; ping. It responds well to nitrogen fertilizers and to Lyon 1984; McLean and Clark 1980; Seidel and others being integrated with legumes (Shaw and Cooper 1990). It should not be included in seeding mixes with 1973). In areas with dry, cold winters, a hardy variety native species.

Varieties and Ecotypes Varieties that have been developed and for which seed is available include: ‘Able’, ‘Akaroa’, ‘Amba’, ‘Am- bassador’, ‘Arctic’, ‘Benchmark’, ‘Cambria’, ‘Chinook’, ‘Comet’, ‘Dactus’, ‘Dart’, ‘Dawn’, ‘Hallmark’, ‘Haymate’, ‘Juno’, ‘Justus’, ‘Kay’, ‘Latar’, ‘Napier’, ‘Pomar’, ‘Potomac’, ‘Prairial’, ‘Rancho’, ‘Rapido’, ‘Sumas’, and ‘Summer Green’ (Alderson and Sharp 1994). The only two rangeland varieties are ‘Berber’ and ‘Paiute’. Berber orchardgrass was developed for California rangelands from material obtained from South Aus- tralia (Alderson and Sharp 1994). It has not been seeded in the Intermountain West. Paiute orchardgrass was introduced from Ankara, Turkey (Alderson and Sharp 1994). It is a drought- tolerant accession that is adapted to basin big sagebrush, pinyon-juniper, mountain brush, mountain big sagebrush, and ponderosa pine communities receiving 14 or more inches (355 mm) of annual precipitation. Paiute is characterized by an abundance of thick, succulent basal leaves that tend to remain green throughout the winter. It begins growth in spring as the snow melts. Plants are palatable and sought out by big game and livestock. Seed should be fall planted on a firm seedbed and covered. Germination is slow, but once established, Paiute forms strong bunches that are very persistent. Plants are shade tolerant and Figure 23—Orchardgrass seedings provide forage, compatible with some seeded and indigenous species. stabilize disturbances, and suppress the spread of Latar, an introduction from Russia, and especially invasive species, but it interferes with the recovery Potomac, developed from collections made in old and establishment of natives (RMRS photo). pastures on the east-central coast of the United States

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(Alderson and Sharp 1994), are used extensively in the Intermountain West for seeding streambanks and disturbed sites in mountain brush, aspen, ponderosa pine, and mountain big sagebrush communities. These two varieties produce an abundance of succulent leaves just after snowmelt in spring and remain green until late fall. Both varieties are long lived and exhibit considerable shade and grazing tolerance. They should generally be seeded in fall, and they can be planted with other rapidly developing species. Germination is generally slow.

Deschampsia caespitosa Tufted Hairgrass ______Synonyms Aira cespitosa Deschampsia pungens Deschampsia holciformis Deschampsia beringenis

Description Tufted hairgrass is a caespitose, mat forming perennial with culms 0.7 to 4.3 ft (2 to 13 dm) tall, and smooth, hairless leaf sheaths. Leaf blades are firm, flat, or folded, 0.04 to 0.20 inch (1 to 5 mm) wide, and often roughened ventrally. Ligules are 0.12 to 0.37 inch (3 to 9.5 mm) long, lacerate along the margins, and gradually tapering to a sharp point. Panicles are 1.6 to 11.8 inches (4 to 30 cm) long, loose, often nodding, and open to narrow, with slender, hairlike Figure 24—Tufted hairgrass exhibits considerable branches that are fine hairy. Spikelets are two or three ecotypic variability; individual populations are adapted flowered and usually shiny and purplish. Glumes are to a wide variety of soil types (Hitchcock 1950). lance shaped, acute, and smooth or roughened. The first glume is single nerved and 0.08 to 0.20 inch (2 to 5 mm) long. The second glume is 0.10 to 0.20 inch (2.5 to 5.2 mm) long and one to three nerved. Lemmas Ecological Relationships and Distribution are 0.10 to 0.18 inch (2.5 to 4.5 mm) long, often purplish basally, and five nerved with the four lateral A circumboreal species, tufted hairgrass is distrib- nerves ending in apical teeth. The awns are 0.08 to uted in the Western Hemisphere from Alaska to New- 2.5 inches (2 to 6.5 cm) long and attach below mid- foundland, south through the Western and Eastern length on the lemma. The callus is densely hairy. States and into Northern Mexico. In the more south- Paleas more or less equal the lemmas. Plants flower erly portions of its range it grows at higher elevations. from late July to September (Arnow 1987, 1993; Its distribution in the Great Plains is limited (Hitchcock Cronquist and others 1977; Hitchcock 1950; Hitchcock 1950; Larson 1993; Stubbendieck and others 1992). and others 1969; USDA Forest Service 1937). Tufted hairgrass occurs in coastal prairies, bogs, Tufted hairgrass has been described as 30 or more marshes, wet to mesic meadows, forest openings, dry different races, variants, or species. Recent western slopes, aspen stands, high-elevation valley bottoms, floras have included these in the highly variable and open areas above timberline, particularly on sites species D. caespitosa (see Cronquist and others 1977) where snow accumulates (Gould and Shaw 1983; (fig. 24). Johnson and Billings 1962; Ward 1969). It is also found along ditches, streams, and lakeshores.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 369 Chapter 18 Grasses

Tufted hairgrass is a long-lived, cool-season species viability of wildland collections are highly variable that exhibits considerable ecotypic variation (Ward (Chambers 1989). Germination requirements may vary 1969). It occurs at elevations from sea level to over among seed lots. Archibald and Feigner (1995) re- 14,000 ft (4,270 m) (Brown and others 1988) that ported that seeds do not require moist prechilling for receive 14 to 24 inches (356 to 609 mm) of annual germination. Other investigators, however, found that precipitation (USDA Natural Resources Conservation exposure to light or transfer from a warm (68/50 ∞F) (20/ Services 2002). It is more common on sites that receive 10 ∞C) (16 hrs/8 hrs) environment to a cooler environ- at least 20 inches (508 mm) of annual precipitation ment (59/41 ∞F) (15/5 ∞C) (16/8 hrs) after 6 weeks of (Brown and others 1988). Tufted hairgrass is adapted incubation improved germination of their seed lots to fine- to medium-textured neutral to somewhat acidic (Chambers and others 1987; Guerrant and Raven soils that are of moderate depth and fertility. Some 1995; Sayers and Ward 1966). populations occur on coarser sandy or gravelly soils Seed fields are planted at a rate of 1 pound per acre (Gates 1962; Johnson 1962; Lewis 1970). Plants are (1.1 kg per ha). Irrigated fields may be planted in tolerant of anaerobic conditions (USDA Natural Re- spring or fall. Seeds are very small and should be sources Conservation Services 2002). They are not drilled no more than 0.25 inch (0.6 cm) deep. Weeds drought tolerant, but they exhibit good winter hardi- must be controlled, as the early growth rate of tufted ness (USDA Natural Resources Conservation Service hairgrass seedlings is only moderate. Mature stands 2002). Individual populations tolerant of salinity, al- are quite competitive. Seed yields average 40 pounds kalinity, or specific heavy metals have been identified per acre (44.8 kg per ha) (Smith and Smith 1997). (Frenkel and Morlan 1991; Hardy BBT Limited 1989; Wildland seedings are conducted in late fall (Brown Rabe and others 1999). and others 1988; Chambers and others 1987) or in Tufted hairgrass occurs in a large number of late spring at higher elevations. Seeds can be drilled or successional or climax communities (Walsh 1995), broadcast and covered at 0.8 to 1.6 pounds PLS per growing mixed with other species or as nearly pure acre (0.9 to 1.8 kg per ha) or greater on highly dis- stands on open sites, particularly at higher elevations. turbed sites (Pawnee Buttes Seed, Inc. 2002). It rarely occurs as an understory species in shaded Plugs, sod, and container stock have also been used areas (Chambers 1987). Tufted hairgrass is also a to establish tufted hairgrass on disturbances and common species on disturbed areas within its range, provide soil stabilization more rapidly. Applications of appearing in early- to midsuccessional communities fertilizers and soil amendments aid in hastening es- (Brown and others 1988). tablishment on infertile soils (Brown and Chambers Plants begin growth in early spring when tempera- 1989, 1990; Brown and others 1988). tures are near freezing. In moist areas the leaves Tufted hairgrass is a valuable revegetation species, remain green throughout the growing season (Bell particularly for disturbances at high elevations 1974; Lewis 1970; Vallentine 1961). New leaf growth (Vaartnou 1988). Because of the considerable ecotypic is initiated prior to the onset of dormancy, and imma- variability, local seed sources or sources known to be ture leaves are protected over winter by dry leaves of adapted to the planting site should be used. This is the previous season. particularly true on sites with unusual soil character- Tufted hairgrass is fairly fire tolerant, as the leaves istics. Populations have been identified that are toler- are generally green during the growing season and ant of heavy metals such as copper, lead, and manga- protect the root crown (Great Plains Flora Association nese; other populations will grow on peat, while still 1986). Stands recover through resprouting from the others have been found colonizing coal mine spoils root crown or through establishment of new seedlings (Hardy BBT Limited 1989; Russell 1985; Vitt and from the soil seed bank (Chambers 1993; Gehring and others 1990). Some populations are adapted to rela- Linhart 1992). Severe fires can kill the plants tively dry sites (Arnow 1987). (DeBeneditti and Parsons 1979, 1984). Uses and Management Plant Culture Tufted hairgrass provides an abundant source of Tufted hairgrass is self-sterile. Fruits ripen in sum- forage from spring through summer. It provides good- mer. Wildland stands may be hand harvested in sum- quality summer range and is one of the most valuable mer, but early shattering can reduce seed yields. forage species at high elevations. Tufted hairgrass Combining or swathing followed by combining are meadows are sometimes cut for hay. Late summer and options for harvesting cultivated seed fields and ex- fall regrowth are moderate. tensive wildland stands on level terrain. Harvested Although fairly tolerant of close grazing, the species seed is cleaned using air screen machines and a does decrease with excessive grazing and associated debearder or huller-scarifier. There are about 2,500,000 trampling. It is considered an indicator species for seeds per pound (5,500,000 seeds per kg). Fill and grazing pressure in many high-elevation communities.

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Stands should be allowed to set seed periodically to perpetuate the community. Management activities that alter the water table can also impact tufted hairgrass. Increased shading with development of woody species in later successional communities reduces tufted hairgrass populations. Tufted hairgrass is used to seed mined lands, dredge spoils, coal mine overburden, burned forest lands, degraded livestock and game ranges, ski areas, and other disturbances at high elevations (McVay and others 1980; Pawnee Buttes Seed, Inc. 2002). Seed sources adapted to local soil conditions and with appropriate cold tolerance should be used for these projects.

Varieties and Ecotypes ‘Norcoast’ was selected from collections made in Cook Inlet, Alaska (Alderson and Sharp 1994), and released for revegetation of disturbances and to provide forage. It has been used on adapted sites from Alaska to California, but it has not seen use in the Figure 25—A strongly rhizoma- Intermountain West. tous warm-season perennial, in- ‘Nortran’ was developed from materials collected in land saltgrass can grow on sites Alaska and Iceland (Alderson and Sharp 1994; Mitchell with shallow water tables and on 1988). It is used primarily for the revegetation of highly saline soils (photo cour- disturbed areas and for pasture seedings. It is hardy in tesy of Marshall Haferkamp, alpine and boreal regions south of the Arctic. It has not USDA ARS, Miles City, MT). been seeded in the Intermountain West. ‘Peru Creek’ was developed from mass collections made in subalpine meadows along Peru Creek, near Dillon, CO (Englert and others 2002), at an elevation scaly rhizomes at shallow depths, generally in the of about 10,200 ft (3,120 m). It has been tested and upper 8 inches (20 cm) of the soil profile. The culms seeded in the Intermountain West for reclamation of sometimes bear long, soft, straight hairs that are mined lands and other disturbed sites with acidic soils about 0.02 to 0.12 inch (0.5 to 3 mm) long at the collar and for restoration of wet meadows at high elevations. and upper margins of the sheath. Leaf blades on the middle or upper portions of the culm are two ranked, Distichlis spicata loosely involute and 0.08 to 0.16 inch (2 to 4 mm) wide, with the upper surface sometimes roughened or long Inland Saltgrass ______hairy. Ligules are 0.01 to 0.02 inch (0.2 to 0.6 mm) long with a fringe of cilia along the margins. Panicles are Synonyms 1.2 to 4.0 inches (3 to 10 cm) long with a few somewhat Uniola spicata congested and strongly compressed spikelets. The Uniola stricta male panicle often extends above the leaves, while the Distichlis maritima var. laxa female panicle is normally enclosed within them. Stami- Distichlis dentata nate spikelets are 6 to 14 flowered and yellowish, while Briza spicata the pistillate spikelets are 3 to 10 flowered. Glumes are lance shaped, hairless, and five to seven nerved. Description Lemmas are firmer than the glumes and seven to nine nerved. Glumes and lemmas of pistillate flowers are Inland saltgrass is a native, dioecious, warm-sea- larger than those of the staminate flowers. The paleas son, strongly rhizomatous perennial halophyte (Ungar of the pistillate florets are slightly shorter than the 1974) that is widely distributed across the Western lemma, and their winged keels have irregular, shal- United States and Canada (fig. 25) (Hitchcock 1950). lowly serrate, and minutely ciliolate margins (Arnow It is a relatively short grass that grows to 16 inches 1987; Cronquist and others 1977; Hitchcock 1950; (40 cm) in height. Inland saltgrass produces vigorous Hitchcock and others 1969).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 371 Chapter 18 Grasses

Saltgrass is best adapted to medium- and fine- tolerance of flooding (Platts and others 1987). Al- textured soils (Shaw and Cooper 1973), but it can also though flooded plants may show a loss of vigor, they be found in sandy and gravelly soils. It often occurs in generally recover within 30 days after the water has conjunction with a shallow water table and on sites receded. Saltgrass tolerates burning, as the rhizomes that are periodically flooded. On highly saline soils, are generally insulated from the heat of most fires. plants may be dwarfed, and the culms are generally reclining with the tips ascending, or they may be Plant Culture decumbent at the base and somewhat stoloniferous. Saltgrass has a number of physiological adaptations Inland saltgrass reproduction is mainly through that allow it to occupy saline soils. Vesicular-abuscular vegetative spread from a well-developed system of mycorrhizal fungi have been observed on inland deep underground rhizomes. It is well adapted to saltgrass roots, and are thought to enhance salt toler- spreading into a wide variety of soils. The species is a ance (Ho 1987). Plants contain salt glands that are poor seed producer. Seed production is generally re- active in the extrusion of salt (Hansen and others stricted to better sites that support dense healthy 1976). The lacunae tissue of the roots is apparently stands (Cluff and Roundy 1988). Fill and viability of continuous with the rhizome and leaf sheath. This saltgrass seed is often quite high. Seed can remain allows for gas exchange under partial inundation and dormant for at least 4 years (Shaw and Cooper 1973). in heavy soils (Hansen and others 1976). This allows soil seed banking to occur and permits plants to establish in large numbers when favorable Ecological Relationships and Distribution conditions exist for germination. Saltgrass seed requires relatively high temperatures, low salinities, Inland saltgrass is distributed from the interior of and moist soil to germinate (Roundy 1987). Saltgrass British Columbia to Saskatchewan and southward to has been successfully seeded with a Brillion seeder California, Texas, and Mexico. It also ranges eastward and with broadcast seeding. Seed must be planted to Minnesota. Inland saltgrass is dominant or codomi- about 0.5 inch (1.3 cm) deep. nant in some communities and may be found in vary- Saltgrass has been established by planting rhi- ing densities in a wide range of communities. Pure to zomes. Because rhizomes are sensitive to desiccation, nearly pure stands occur on sites where soils are saline they should be planted only in wet, saline areas, unless and fine textured, and where the soil is wet at least irrigation is being used during establishment. Plant- part of the year (Shantz and Piemeisel 1940; Ungar ing rhizomes at depths of 1 to 2 inches (2.5 to 5 cm) has 1974). In the Great Basin saltgrass is a dominant proven successful (Delzell 1972). species in lower elevation desert shrub communities. It spreads rhizomatously and forms a sod under black Uses and Management greasewood and shadscale. Black greasewood and saltgrass occurring together indicate the presence of Saltgrass is not greatly preferred by livestock and soil with a high moisture content. Where soil salinity wildlife. Cattle generally avoid its harsh foliage. Be- becomes greater than 1 percent, black greasewood cause it usually remains green all summer long, it is gives way to pure stands of saltgrass (Brotherson most often grazed after other grasses have been con- 1987). sumed or have dried out (Humphrey 1970; Shaw and Saltgrass does especially well on fine-textured soils Cooper 1973). Saltgrass is low in nutritive value. For in areas with high water tables that receive seasonal cattle, a pure saltgrass diet should be avoided because runoff. Salt flats, desert playas, valley bottoms, inter- severe rumen compaction can occur (Stubbendeick mittent ponds, saline meadows, and borders of springs, and others 1985). streams, and lakes commonly support this species. Small mammals and birds use saltgrass for cover, Inland saltgrass forms a dense sod only on salt soils nesting, and feeding. It often forms a dense sod that with fine texture and good soil moisture throughout provides good escape and security cover. Saltgrass most the year. It may also occur as a pioneering stands can provide nesting cover for pheasants and species. The spreading rhizomes allow saltgrass to many species of ducks and shore birds. Seeds of colonize salt flats, playas, lakeshores, shale cliffs, and saltgrass are eaten by waterfowl and small mammals other barren saline sites (Cronquist and others 1977; (Ohlenbusch and others 1983). Saltgrass sod around Unger 1966) not tolerated by other grasses. Its pointed lakes often indicates alkaline conditions that are un- rhizomes are well adapted to heavy clays and shales favorable for fish (Hansen and others 1988a,b). (Hansen and others 1976). Saltgrass is highly resistant to grazing and tram- Saltgrass has moderate to high drought tolerance pling, and it provides a resistant sod in areas where and can remain dormant for long periods of time cattle concentrate, such as watering sites, corrals, and (Shantz and Piemeisel 1924). It exhibits exceptional along trails. Attempts have been made to rehabilitate saltgrass meadows by killing the saltgrass and planting

372 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses more palatable, salt-tolerant species (Bowman and tapering into a long, rough, flexible, sharp awn that others 1985; Ludwig and McGinnies 1978). Plowing, measures 0.63 to 1.18 inches (16 to 30 mm) in length. disking, and other mechanical treatments have proven Flowering occurs from late June through August ineffective because inland saltgrass reproduces pro- (Arnow 1987, 1993; Cronquist and others 1977; fusely from rhizomes (McGinnies 1974, 1975). Hitchcock 1950; Hitchcock and others 1969). E. canadensis has been found to hybridize with Plant Culture Agropyron trachycaulum and E. interruptus (Arnow 1987; Great Plains Flora Association 1986). It has Reestablishment of saltgrass is often required to formed synthetic hybrids with Sitanion hystrix and restore depleted wetlands. Seed, however, is not widely some Agropyron species (Dewey 1967). available. In addition, removal of weedy competition and preparation of a seedbed can be difficult on inland Ecological Relationships and Distribution saltgrass sites. Seeding on sites with some surface litter is advised because the soils often crust, reducing Canada wildrye is distributed from Alaska south to seedling emergence. Inland saltgrass develops slowly, California, and eastward across much of Canada and requiring 2 to 3 years to become fully established. the United States except for the extreme Southeastern Seeding pure stands is normally recommended. United States (Great Plains Flora Association 1986). It generally occurs in areas receiving 20 to 45 inches Varieties and Ecotypes (508 to 1143 mm) of annual precipitation. The species occurs widely on tall-grass and mixed-grass prairies of There are no releases of inland saltgrass. the Great Plains and Midwestern States, particularly in sandy areas (Barker and others 1985). It also grows Elymus canadensis in riparian areas and wetlands, occurring with eastern cottonwood, green ash, redosier dogwood, and Canada Wildrye ______other riparian species in the Missouri River system of Synonyms the Great Plains (Boggs 1984). Canada wildrye is tolerant of shade and occurs in oak savannas of Min- Hordeum patulum nesota (Tolstead 1942), ponderosa pine communities Clinelymus canadensis in the Southwest (Lavin 1953), and mixed evergreen Sitanion brodiei and hardwood forests of the Coast Range in California Elymus glaucifolius (Hickman 1993). It is less common in the Intermoun- Elymus robustus tain West, usually growing in low, moist valleys or Elymus brachystachys along streams, irrigation ditches, and roadsides. Canada wildrye is adapted to soils of low to moder- Description ate fertility. It can be found on soils ranging from gravelly to clayey and from neutral to slightly alkaline Canada wildrye is a highly variable, perennial, cool- (Great Plains Flora Association 1986). Pure stands season bunchgrass. Plants are coarse and open with can be found on wet to moist, alkaline, or sometimes erect to decumbent, tufted, hollow culms 2.6 to 6.5 ft (8 saline meadows (Arnow 1987). to 20 dm) tall. Rhizomes are short and usually absent. Canada wildrye develops rapidly and is rather short Leaves are mostly cauline. The blades are flat, 0.12 to lived. It is a good seed producer and depends upon seed 0.59 inch (3 to 15 mm) wide, and roughened above. for spread and persistence (Weaver and Zink 1946). Auricles are well developed, clasping, and about 0.04 The species is considered early to midseral and often to 0.08 inch (1 to 2 mm) long. Ligules are 0.01 to 0.08 occurs on riparian and upland disturbances. Canada inch (0.2 to 2 mm) long, truncate, and fringed with wildrye begins growth in early spring, flowers in minutely ciliate hairs. Spikes are thick, bristly, and midsummer, and matures seed in late summer or usually nodding or drooping (Asay and Jensen 1996b). early fall. Plants may enter dormancy in summer and Spikelets are 0.47 to 0.59 inch (12 to 15 mm) long resume growth in fall if moisture is received. excluding the awns, two to four flowered, and two to Fire may top-kill Canada wildrye. Its buds are four per node. Flowers are widely separated, leaving located beneath the soil surface where they are some- the axis of the spikelet visible. Glumes are subequal, what protected from burning. The coarse leaves and narrowly lanceolate, about 0.39 to 0.98 inch (10 to stems are not highly flammable. Early spring fires are 25 mm) long, three to five nerved, broadest above the most harmful, while the best postburn response fol- base, and tapering to a long awn. The nerves on the lows summer fires. Regeneration following fire occurs glumes are roughened to somewhat ciliate. Lemmas through recruitment from the seed bank (Howe 1994; are about 0.33 to 0.55 inch (8.5 to 14 mm) long, Mowat 1990; Robocker and Miller 1955; Wright 1971; strongly roughened, five to seven nerved, and gradually Wright and Bailey 1982).

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Plant Culture 1983). It may also be used alone or in mixtures to seed disturbances in sand dunes, riparian areas or wet- Canada wildrye may outcross or self-fertilize (Gabel lands, or native prairie communities. Canada wildrye 1984). It produces numerous reproductive stems and provides good cover for birds and many other small is generally a good seed producer. Seed maturation animals. Its foliage tends to be rather coarse and not occurs from July in the southern portion of the species highly palatable. range to August in the northern portion (Smith and Smith 1997; Wheeler and Hill 1957). Seed production Varieties and Ecotypes is generally good, and shatter during harvest is generally not problematic. However, threshing and cleaning Mandan was released in 1946 by the Agricultural are complicated by the presence of the long awns Research Station Northern Great Plains Research (Hafenrichter and others 1949). Swathing or wind- Center in Mandan, ND, the Natural Resources Con- rowing during the hard-dough stage and combining servation Service and the North Dakota Agricultural are the recommended methods of harvest (Smith and Experiment Station. It originated from material col- Smith 1997). Seed is cleaned using a using a debearder, lected near Mandan, ND, and was selected for its rapid a four-screen fanning mill, and a two-disc indent establishment, leafiness, productivity in young stands cleaner (Smith and Smith 1997). Seeds average 115,000 of mixed species, and soil stabilizing capabilities seeds per pound (254,000 per kg). Seed yields from (Alderson and Sharp 1994; Baker and others 1985). It nonirrigated fields range from 150 to 300 pounds per is shorter and more persistent than many Canada acre (168 to 336 kg per ha), while yields from irrigated wildrye populations. Mandan currently receives little fields range from 400 to 1,100 pounds per acre (448 to use, and seed is generally not available. 1,232 kg per ha) (Smith and Smith 1997). Appropriate seed sources for proposed planting sites should be selected due to the considerable ecotypic Elymus cinereus variation within the species. Seed should be planted in Great Basin Wildrye ______late fall or early spring at a depth of about 1 inch (2.5 cm) by broadcasting and covering the seed or by Synonyms drilling. Seeds germinate rapidly. Dry storage or a Leymus cinereus 2-month moist prechill improve germination (Blake Elymus condensatus var. pubens 1935; Greene and Curtis 1950). Seedlings are vigorous Elymus piperi and stands develop rapidly. Peak production occurs in 2 or 3 years (Wasser 1982). Stand life for seed fields is Description 7 or 8 years (Smith and Smith 1997). In native stands Canada wildrye may eventually be replaced by longer Great Basin wildrye is a coarse, robust, densely lived species. It is not competitive with aggressive tufted perennial bunchgrass that often forms clumps exotic grasses. Canada wildrye is susceptible to infes- up to 3.3 ft (1 m) across. It is the largest cool-season tation by ergot, which may cause illness or death if bunchgrass native to the Western United States consumed by livestock (Vallentine 1961). (fig. 26) (Abbot and others 1991; Anderson and others 1995; Cash and others 1998). Plants sometimes pro- Uses and Management duce short, stout rhizomes. Culms are usually more than 2.3 ft (7 dm) tall, often growing to heights of 6.6 Canada wildrye is used by wildlife and livestock. ft (2 m). They are hairless, or more often minutely Forage value and palatability are greatest in early hairy, especially at the nodes. Leaf sheaths are hair- spring, but decrease as plants mature (Hoover and less to soft hairy with appressed or spreading hairs. others 1948; Morris and others 1950; Stubbendieck Auricles are well developed to lacking. Ligules are and others 1986). Stands of Canada wildrye are har- long and membranous or translucent, measuring about vested for hay in the boot stage, just as the seedheads 0.08 to 0.28 inch (2 to 7 mm) in length. Leaf blades are emerge (Wheeler and Hill 1957). Plants tend to be flat, or nearly so, 0.17 to 0.59 inch (4.5 to 15 mm) wide, avoided after this stage. Stands decrease with heavy firm, and sometimes strongly nerved. Spikes are 2.8 to grazing because the growing points are damaged 9.0 inches (7 to 23 cm) long, 2.8 to 4.7 inches (7 to 12 cm) (Ehrenreich and Aikman 1963). wide, stiff, erect, and sometimes compound and branch- Canada wildrye seedings provide cover, litter, and ing. Spikelets are 0.35 to 0.79 inch (9 to 20 mm) long organic matter; control erosion; and provide habitat and two to five flowered. There are usually three to six for birds and other wildlife (Barker and others 1985; per node, but sometimes only two. Glumes are subequal, Noyd and others 1995). It is tolerant of heavy metals, 0.28 to 0.63 inch (7 to 16 mm) long, narrow, somewhat and can be used to revegetate abandoned mine tailings awl shaped, tapered, and often as long as the spikelet. (Chambers and Sidle 1991; Hardell and Morrison

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prairies, hillslopes, and mountainous sites. Great Basin wildrye invades disturbed sites, and is an important colonizer of steep, eroded slopes and gullies where it is adapted. Great Basin wildrye exhibits considerable ecotypic variation with populations growing at elevations ranging from 1,970 to 9,840 ft (600 to 3,000 m) on sites receiving 8 to 20 inches (200 to 500 mm) of annual precipitation. Some populations grow on sites receiving as little as 5 inches (130 mm) of annual precipitation (Cash and others 1998; Davenport Seed Corpora- tion 1997; Hickman 1993). Great Basin wildrye commonly occurs on deep, silty to clayey soils, but it also occurs on soils that are sandy or gravelly or that have claypan layers or high potassium concentrations. Figure 26—Great Basin wildrye provides good win- Various ecotypes are moderately tolerant of acid, alka- ter cover for birds and small mammals (photo cour- line, or saline conditions (Asay 1987; Wasser 1982). tesy of John Kinney, USDA FS, Rocky Mountain Research Station, Boise, ID). Great Basin wildrye plants begin growth in early spring (Hafenrichter and others 1968). Flowering occurs in early to midspring, and seeds ripen in late summer or early fall (Arnow 1987; Meyer and others 1995). Plants tolerate water stress; growth continues Lemmas are 0.28 to 0.47 inch (7 to 12 mm) long; into summer and sometimes after seed ripening (Ander- pubescent with very short, stiff hairs or rarely hair- son and others 1987). Some ecotypes are moderately less; and usually nerveless below and five to seven tolerant of intermittent high water tables, moderate nerved above. The awn is short, usually less than 0.20 soil drainage, and winter or spring flooding (Barkworth inch (5 mm) in length, or absent. Flowering occurs and Atkins 1984; Lesperance and others 1978; Morris from June to early August (Arnow 1987, 1993; and others 1950). Reproduction is from seed or by Cronquist and others 1977; Hitchcock 1950; Hitchcock tillering or sprouting from rhizomes. and others 1969). Great Basin wildrye plants in the Plants are resistant to fires, particularly those occur- Western United States are generally tetraploid with ring late in the growing season following the onset of 2n = 4x = 28, while those in Canada are octoploid with dormancy (Range and others 1982; Zschaechner 1985). 2n = 8x = 56 (Asay and Jensen 1990b). Great Basin The bunching habit and the presence of coarse, dried wildrye forms natural fertile hybrids with beardless leaves covering the growing points in late summer wildrye and bottlebrush squirreltail in areas where provide protection if burns are not severe (Range and the species overlap (Arnow 1987; Asay and Jensen others 1982; Zschaechner 1985). Damage is greater 1996b). earlier in the season, particularly if soils are dry, limiting the potential for resprouting from the root Ecological Relationships and Distribution crown or rhizomes (Bradley and others 1992a,b; Wasser 1982). If moisture conditions are favorable, production Great Basin wildrye occurs throughout the Western may increase and remain high for several years follow- United States and Canada and reportedly as far east ing fire (Klebenow and others 1976). In addition to as Minnesota (Arnow 1987; Asay and Jensen 1996b). resprouting following fire, stand increase may also In the Intermountain West it occurs in a wide range of occur from the seed bank, as smoke has been found to community types from salt desert shrublands to pon- increase the growth of germinants (Blank and Young derosa pine communities (Arnow 1987). In this area it 1998). commonly occurs with big sagebrush, rabbitbrush, junipers, and wheatgrasses. In the Great Plains it is Plant Culture associated with grama grass, buffalo grass, and mixed- grass prairie communities (Shiflet 1994). Great Basin Wildland stands of Great Basin wildrye may be wildrye may occur scattered among other species or it hand harvested. Extensive wildland stands and seed may form pure stands or stringers along riparian fields may be swathed and combined. Shattering is areas, dry washes, roadsides, and on floodplains or in only moderate, but lodging may create problems. Seeds other areas that receive runoff water or subirrigation, are cleaned by combining or by chopping and fanning or that have high water tables (Jankovsky-Jones and (Plummer and others 1968). There are about 95,000 to others 1999; Wasser 1982). It also grows on upland 166,000 seeds per pound (210,000 to 366,000 per kg)

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(Vallentine 1971). Seed fill and viability are variable depletes carbohydrates and is detrimental to the plants, and often low. Seeds may be marketed with 85 percent especially when they are grazed during the boot stage purity and 80 percent germination (Wasser 1982). (Tweit and Houston 1980). Plants should not be grazed Great Basin wildrye is usually difficult to establish until they reach heights of 12 to 15 inches (30 to 38 cm), due to low or erratic germination, poor seedling sur- and 20 percent of the production should be left to vival, and slow establishment (Evans and Young 1983; maximize regrowth (Hafenrichter and others 1968). Roundy and others 1983). It should be seeded in late Livestock should be removed before the soil dries out fall in the drier portion of its range. Seeds should be to permit regrowth and replacement of root reserves. drilled or broadcast and covered to a 1-inch (2.5-cm) This approach should also be followed for fall grazing. depth (Wasser 1982). Seeds are nondormant and do Great Basin wildrye is susceptible to infection by not require afterripening, but they will germinate ergot, which may cause sickness or death of livestock more readily in spring following exposure to cool, that consume it (Cronquist and others 1977). Plants moist seedbeds in winter (Meyer 1994). This allows are also sometimes severely infested with leaf rusts. the seed to emerge more uniformly in spring and Great Basin wildrye is seeded to provide forage and maximizes survival in the face of drying seedbeds. cover for livestock and wildlife. It is also planted in High-elevation and spring-flooded sites can be seeded pastures, sometimes with tall wheatgrass, on saline in spring as soon as they become accessible. and alkali soils that are subirrigated or that may flood Great Basin wildrye can be drill or broadcast seeded intermittently to provide fall, winter, or spring live- alone or with other nonagressive species in areas stock grazing and for calving. These seedings are also where it is most likely to establish. It should not be harvested for their hay and silage production. seeded in mixtures with more aggressive species or in Great Basin wildrye is valuable for erosion control areas where weeds have not been controlled. High soil plantings in areas where it is adapted because of its salinity or soil crusting may reduce emergence. Stand deep and spreading fibrous root system. It is useful for establishment requires two or three growing seasons. stabilization of drainage ditches, mine spoils, highway New seedings of Great Basin wildrye should not be rights-of-way, burned sites, eroding slopes, and other grazed or harvested until after the first two growing disturbances (Barker and others 1985; Cronquist and seasons. others 1977; Walker and Brotherson 1982). Great Basin wildrye is also seeded as a component of wind- Uses and Management breaks and conservation plantings. A major use of the species is for seeding disturbances on alkaline and Great Basin wildrye is an important forage species saline sites that once supported the species. Monsen for wildlife and domestic livestock. It is palatable and (1983) recommends that basin wildrye be included in nutritious in spring and fall and receives considerable seedings on disturbances in or adjacent to riparian use during these seasons (Jarecki 1985). It is also areas including wet meadows, stream terraces, and valuable in winter because of its height and the acces- mountain brush, big sagebrush, desert shrub, and sibility of its forage (Cook and others 1954; Jarecki saltgrass communities. Slow establishment, however, 1985). Palatability and nutrient content decrease in limits the value of Great Basin wildrye during the first summer and winter as the leaves dry and become two growing seasons following seeding. coarse (Cook and others 1954; Daubenmire 1970; Jarecki 1985; Krall and others 1971; Walker and Varieties and Ecotypes Brotherson 1982; Wasser 1982). Big game animals including mule deer, elk, and ‘Magnar’, an octoploid cultivar, originated in bighorn sheep graze Great Basin wildrye (Austin and Saskatchewan, Canada (Alderson and Sharp 1994), others 1994; Keating and others 1985; Majerus 1992). and has been seeded extensively throughout the Inter- It provides excellent cover for small mammals, water- mountain West in upper pinyon-juniper, mountain fowl, and upland game birds, and bedding cover for big brush, and mountain big sagebrush communities. It game (Batzli 1974; Sours 1983; Wasser 1982). also establishes well in canyon bottoms, along road All classes of livestock use Great Basin wildrye. It rights-of-way, and in depressions where extra mois- provides valuable forage and thermal protection for ture accumulates. It is adapted to soils ranging from livestock in winter. Native stands provide spring and loams to clays, and from slightly basic to alkaline. fall grazing and may be cut for hay. Poor grazing Magnar performs best on sites receiving 15 inches practices, however, have degraded many native stands. (380 mm) or more of annual precipitation, but stands The timing, frequency, and extent of grazing all have establish in areas receiving as little as 13 inches an impact on plant vigor and regrowth (Holzworth and (330 mm). It is not suited to lower, drier pinyon- Lacey 1991; Jarecki 1985). Forage is of high quality in juniper or sagebrush grass communities. early spring. Excessive grazing during this period

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Magnar foliage is blue green. Plants grow to 7 ft straight awns that are about 0.39 to 1.18 inches (10 to (2.1 m) in height and produce wide, coarse leaves and 30 mm) long. Plants flower from June to August large stems and seed heads. Seedlings are vigorous (Arnow 1987, 1993; Cronquist and others 1977; and establishment is generally rapid. Magnar is long Hitchcock 1950; Hitchcock and others 1969). lived, nonaggressive, and suitable for mixed seedings. Blue wildrye is often confused with Agropyron ‘Trailhead’ was collected near Roundup, MT trachycaulum var. unilaterale. Blue wildrye, however, (Alderson and Sharp 1994). Plants are tetraploid and has the Elymus characteristics of two spikelets per dark green with broad, somewhat lax leaves. Trailhead node, narrow glumes, and a distorted rachilla. It is, is considerably more drought tolerant and productive however, extremely variable in its morphological and than Magnar. It has established successfully in areas genotypic characters. Some biotypes cross readily with receiving as little as 11 inches (280 mm) of annual Sitanion hystrix, while others do not. These hybrids precipitation. Trailhead is adapted to soils ranging are referred to as X elysitanion hansenii. Some manu- from saline-alkaline to acidic. It can be seeded from als recognize blue wildrye varieties such as E. g. var. valley bottoms up through the mountain brush zone. breviaristatus, E. g. var. glaucus, and E. g. var. jepsonii Trailhead is seeded to provide soil stabilization, early (Cronquist and others 1977). The species is spring and winter forage, and wildlife habitat. allotretraploid with 2n = 28 (Dewey 1982).

Elymus glaucus Ecological Relationships and Distribution Blue Wildrye ______Blue wildrye is distributed from Alaska to Califor- nia and into northern Arizona and New Mexico. It is Synonyms found less commonly from Ontario to New York, and south to Arkansas (Arnow 1987, 1993; Cronquist and Elymus sibericus var. glaucus others 1977). Blue wildrye is most abundant in tim- Elymus villosus var. glabriusculus bered areas from sea level to 10,000 ft (3,050 m) (Asay Elymus nitidus and Jensen 1996b). It occurs in shaded or open forests, grasslands, shrublands, and on moist soils of riparian Description areas and wetlands in mountain brush, aspen, ponde- Blue wildrye is a green to blue waxy cool-season rosa pine, spruce-fir, and lodgepole pine communities native perennial bunchgrass that forms small loose to (Arnow 1987; Hitchcock 1950) (fig. 27). dense tufts with only a few culms. The root system is Blue wildrye populations are adapted to well-drained deep and fibrous. Plants sometimes produces short soils with clay loam to sandy loam textures (Hassell stolons. Culms grow from 1.9 to 4.6 ft (6 to 14 dm) in and others 1982) that are moderately acid to neutral height. Leaves are scattered along the culms. Leaf (pH 5.2 to 7). Populations are common on moist sites, sheaths may be glabrous or with downward pointing but some are moderately drought tolerant. The species hairs; the collar is often purple. Leaf blades are flat to is adapted to soils of low fertility, but it does not sometimes slightly involute, 0.16 to 0.47 inch (4 to 12 tolerate shallow soils, and it is moderately sensitive to mm) wide, and mostly roughened to sparsely silky saline soils (Hassell and others 1983). haired, or sometimes hairless below. Auricles are well Plants are short lived, but stands reseed themselves developed, about 0.08 inch (2 mm) long, and clasping. readily (Asay and Jensen 1996b). The species is fa- Ligules are short, 0.01 to 0.04 inch (0.3 to 1 mm) long, vored by disturbances and is often most abundant in squared at the apex, and irregularly lacerate and early seral communities on open, disturbed sites where ciliate along the margins. Spikes are 2.4 to 6.3 inches it may form dense stands (Cole 1982; Hitchcock and (6 to 16 cm) long, excluding the awns, erect, inter- others 1969; USDA Forest Service 1937). Plants also rupted below and overlapping above, or dense through- grow in shaded areas (Plummer and others 1955). out and long exerted. Spikelets are mostly two per Blue wildrye forms small bunches with coarse foli- node, 0.39 to 0.63 inch (10 to 16 mm) long, and two to age that burn rapidly and can survive wildfires. Burned four flowered. Glumes are nearly parallel, concealing plants regenerate by resprouting from the root crown the base of the enclosed florets and tapering to a short (Simmerman and others 1991). New plants establish awn. They are subequal, 0.25 to 0.75 inch (6.5 to 19 mm) from the seed bank, as burns appear to provide favor- long, three to five nerved, narrowly lance shaped, and able microsite conditions for germination (Stickney broadest above the base. The glumes are hardened 1989). Blue wildrye seedlings and cover develop rap- below and membranous above with glabrous, smooth idly, and stands of the species are common on burned to roughened surfaces. Lemmas are 0.33 to 0.55 inch over sites where competition is low. Stands may de- (8.5 to 14 mm) long, five nerved, hairless to roughened cline after the first few years (Brown and DeByle 1989; along the nerves, and tapering to long, slender, mostly Hafenrichter and others 1949, 1968).

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Some seed is commercially available. Libby and Rodrigues (1992), however, suggested that use of commercial seed outside its area of origin could con- taminate local populations. Hassell and others (1983) recommended use of local seed collected or produced from populations growing within 300 miles (483 km) and 1,640 ft (500 m) elevation from the planting site because of the high variability within the species. Blue wildrye can be drill or broadcast seeded on a firm seedbed in fall or early spring. Seed should be covered to a depth of about 0.5 inch (1.3 cm) (Plummer 1943). Establishment and growth are rapid and seedings are persistent.

Uses and Management Although blue wildrye provides fair to good forage for big game and livestock early in the growing season, the foliage is coarse and provides only poor protein content and good energy value (Dittberner and Olsen 1983). The species produces a fairly extensive root system, but plants are not tolerant of heavy grazing (Vallentine 1961). Blue wildrye recovers fairly rapidly when grazing is removed (Sampson 1944). A widespread species, it often provides considerable forage for wildlife. Early spring growth is most readily used; the awned seedheads are avoided (USDA Forest Ser- vice 1937). Blue wildrye establishes rapidly and is a good seed Figure 27—Blue wildrye is a native, shade-tolerant producer (Plummer and others 1968). Consequently, bunchgrass that can be seeded as an understory in it is used for postfire and postlogging seedings because tree and shrub plantings (photo courtesy of Kevin it generally establishes well and can be planted in Jensen, USDA ARS, Logan, UT). mixtures with other natives (Frishchknecht and Plummer 1955). It is a pioneering species and can be seeded to provide cover rapidly, spread from seed, and control erosion on steep slopes, roadsides, and other disturbances (Darris and others 1996; Hafenrichter and others 1949, 1968). Seedings of blue wildrye do not compete excessively with establishing tree seedlings, Plant Culture but the grass does restrict invasion of weedy species (Asay and Jensen 1996b). Blue wildrye has also been Blue wildrye tends to be a good seed producer. used to provide an understory cover in windbreak and Shattering can reduce yields, but seed can be safely conservation plantings. harvested shortly before reaching maturity and allowed to ripen while drying (Link 1993). Seed must be Varieties and Ecotypes dried before hammermilling to facilitate removal of the awns (Hafenrichter and others 1949, 1968; Link Blue wildrye contains considerable intraspecific vari- 1993). Awns of seeds in large lots may be detached ability manifested by differing site requirements. with a debearder or brush machine. Debris is then Plants are highly self-fertile. Some ecotypes are cross- removed using an air screen machine. sterile, while others are compatible (Asay and Jensen There are about 131,000 seeds per pound (288,800 1996b; Snyder 1950). Seed transfer zones for the per kg) (Hafenrichter and others 1968). Seeds main- species have been identified for National Forests of tain viability for 2 to 4 years. They do not require north-central Oregon. a pretreatment for germination (McLean 1967). ‘Arlington’ originated in Snohomish County, WA Seeds should be incubated at 68/86 ∞F (20/30 ∞C) (Alderson and Sharp 1994). It has been seeded prima- (16 hrs/8 hrs) for germination testing (Haferkamp rily for soil stabilization and to provide forage. It grows and McSwain 1951). in shaded areas under the canopies of aspen groves

378 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses and conifer forests, but it also does well on the forest edge and in openings, especially on moist sites and along stream edges. Arlington exhibits good seedling vigor, establishes readily, and provides cover for erosion control on burned and cutover forest areas. Plants are short lived, but the variety does spread from seed. Seed may be drilled shallowly or broadcast following a fire or on snow in winter. Seed is now rarely available.

Elymus junceus Russian Wildrye ______Synonyms Elymus cretaceus Psathyrostachys juncea

Description

Russian wildrye is a densely tufted, perennial, cool- Figure 28—Russian wildrye is drought tolerant and season bunchgrass with the leaves mostly basal. Culms produces palatable forage throughout the summer are 1.3 to 3.6 ft (4 to 11 dm) tall. Leaf sheaths are (RMRS photo). hairless; the auricles and collars are well developed. Blades are flat, waxy, roughened, involute, and 0.06 to 0.16 inch (1.5 to 4 mm) wide. Ligules are very short, 0.01 to 0.04 inch (0.2 to 1 mm) long, squared at the provides nutritious and palatable forage nearly year apex, and ciliate along the margins. Spikes are erect, long on arid and semiarid rangelands (Plummer and 1.6 to 5.1 inches (4 to 13 cm) long, and 0.16 to 0.47 inch others 1968). It is now distributed from Canada south (4 to 12 mm) wide with the rachis disarticulating to Arizona, and eastward to Nebraska (Arnow 1987). between the spikelets. Spikelets are three or sometimes Scattered naturalized plants have been found in Utah, two per node. They are 0.31 to 0.39 inch (8 to 10 mm) Wyoming, Saskatchewan, and Manitoba (Cronquist long and two to three flowered. Glumes are subequal, and others 1977). In the Intermountain area the spe- short, 0.12 to 0.27 inch (3 to 7 mm) long, awl shaped, cies is adapted to salt desert shrub, sagebrush, grass- roughened or short haired, but often hairless and lands, pinyon-juniper, oak scrub, mountain brush, shiny at the base. Lemmas are small, 0.26 to 0.39 inch aspen, and ponderosa pine communities. It grows in (6.7 to 10 mm) long, roughened to stiff haired, five to openings and in the shade of shrubs, but it is not seven nerved, and tapering to an awn that is 0.02 to adapted as an understory species in the shade of 0.12 inch (0.5 to 3 mm) long (Arnow 1987, 1993; Gambel oak and mountain brush (Plummer and oth- Cronquist and others 1977; Hitchcock 1950; Hitchcock ers 1968). Russian wildrye is adapted to fertile loam and others 1969). and clay soils, but it will grow on a variety of soil types Russian wildrye is easily identified by the readily including moderately saline or alkaline soils. It can be disarticulating axis of the rachis at maturity. This seeded on soils that are too alkaline for crested wheat- results in the accumulation of old leaf sheaths at the grass. Russian wildrye is more heat tolerant than base of the plant that eventually separate into fibers. crested wheatgrass in areas receiving summer pre- Plants flower from May to July (Arnow 1987, 1993; cipitation, and matures later, providing a longer pe- Cronquist and others 1977; Goodrich 1986; Hitchcock riod of grazing. It is best adapted to areas with winter 1950; Hitchcock and others 1969). and summer precipitation or a predominance of summer precipitation. The species is drought and cold Ecological Relationships and Distribution tolerant. It does not tolerate spring flooding (Jensen and others 2001). Russian wildrye, a native of the steppes and deserts of Russia and China, was introduced to the United Plant Culture States in 1927. It has been seeded on the Northern Great Plains and the Intermountain and Rocky Moun- Russian wildrye seed matures in July to August. tain areas as a pasture grass and erosion control Seed production is generally low. Crops can be lost to species since the 1950s (fig. 28). Russian wildrye wind or rainstorms occurring during the ripening

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 379 Chapter 18 Grasses period, and seeds tend to shatter shortly after reach- Varieties and Ecotypes ing maturity. Fields are windrowed or direct combined. They can be grazed following harvest to remove ‘Bozoisky-select’, an introduction from Russia litter (Jensen and others 2001). There are about 170,100 (Alderson and Sharp 1994), was selected for its large Russian wildrye seeds per pound (375,000 per kg) seed size, good seedling vigor and establishment, and (Heady 1975). Fresh or dormant seed becomes more enhanced forage production. It is seeded throughout the Intermountain West in the same areas as ‘Vinall’, germinable following a 5-day prechill at 41 or 50 ∞F (5 but exhibits better establishment and productivity. or 10 ∞C) (AOSA 1999). Seed is incubated at an alter- ‘Cabree’ was developed from Russian material for nating temperature regime of 68/86 ∞F (20/30 ∞C) (16 hrs/8 hrs) for 14 days for germination testing. use on the Northern Great Plains and Canadian prai- Wildland seedings can be established in areas re- ries (Alderson and Sharp 1994). It was selected for its ceiving as little as 8 inches (20 cm) of annual precipi- seedling vigor, forage and seed production, and moder- tation. Seed should be drill seeded or broadcast and ate drought tolerance. It has not been seeded exten- covered in fall at a depth of about 0.75 inch (1.9 cm) sively in the Intermountain West. (Ludwick 1976). Seeds are easily handled and planted. ‘Mankota’, introduced from Alma-Ata in the former Seedling vigor is low to moderate, and slow establish- U.S.S.R. (Alderson and Sharp 1994), was developed ment can preclude stand success when weedy compe- for use on the Northern Great Plains and has received tition is present and during dry periods because few little use in the Intermountain West. It is similar to seminal roots are produced (Jensen and others 2001). Vinall and Bozoisky-select. Stands are seeded at wide row spacings, as established ‘Mayak’ was selected for use on the Northern Great plants are strongly competitive and self-thinning un- Plains and Canadian prairies (Alderson and Sharp til widely spaced. The species is difficult to establish in 1994), and has only infrequently been seeded in the mixes with species having more rapidly developing Intermountain West. Mayak provides excellent forage seedlings. throughout the grazing season and has good curing qualities that make it especially useful for late sum- Uses and Management mer, fall, and early winter grazing. Its characteristics include good forage and seed yield and resistance to Russian wildrye is seeded primarily to provide for- leaf spot disease. age in dry areas because of its basal leaf production. ‘Swift’, derived from ‘Saki’ and ‘Mandan 1546’ Grazing of newly seeded stands should be delayed (Alderson and Sharp 1994), is adapted to areas of the until plants are mature. Established stands are long Intermountain West, even though it was developed for lived and tolerant of grazing on fertile sites; stands the Northern Great Plains and Canadian prairies. It decline more rapidly on sites of low fertility. The was selected for its seedling vigor, early spring growth, species begins growth early in the season, and the forage and seed production, and leafiness. It has not numerous basal leaves develop rapidly. Unlike many proven superior to Bozoisky-select. cool-season grasses of dry areas, it is palatable and ‘Tetracan’ is a colchicine-induced tetraploid (Alderson nutritious from spring into fall and exhibits good and Sharp 1994) with large seedheads and seeds, wide regrowth following use (Jensen and others 2001). leaves, and good seedling vigor. It has performed well Seedings of the species provide good quality and quan- on adapted sites, especially with regard to seedling tities of fall and winter forage when other usable vigor and establishment. Tetracan has seen little use forage is often not available. Although seedlings of this in the Intermountain West. species develop slowly, established stands are more Vinall was introduced from Russia (Alderson and competitive than associated native species. Sharp 1994) and has been seeded widely throughout Russian wildrye is seeded in pastures and main- the Intermountain West, especially in pinyon-juni- tained with fertilizer treatments in some areas (Jensen per, basin big sagebrush, Wyoming big sagebrush, and others 2001). Legumes may be added to improve black greasewood, and salt desert shrub communi- productivity. The species is not generally used for hay ties. It is adapted to clay and clay-loam soils, and it is production. Wildlife use the species, and it can provide moderately tolerant of saline and sodic soils. A major valuable forage in early spring and also in late sum- drawback of this variety is its slow and erratic germi- mer when other grasses have dried out (Dittberner nation and establishment. It does not do well when and Olson 1983; Plummer and others 1968). It pro- seeded with species that establish and develop more vides some cover for small mammals and birds. Be- rapidly. Mature stands can prevent the establish- cause plants tend to be widely spaced, it is not often ment of other species. This variety is long lived and used to control erosion; however, it has been used on drought tolerant. roadside areas in southern Idaho to reduce blowing sand and dust.

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Elymus salinus of alkalinity (Cronquist and others 1977). The species grows on hillsides, alluvial fans, plateaus, bluffs, can- Salina Wildrye ______yons, and mountainous areas (Baker and Kennedy Synonyms 1985; Vallentine 1989) (fig. 29). Plants are generally scattered, and normally do not form pure stands. The Elymus ambiguus var. salina species may be resistant to wildfire due to its rhizoma- Elymus ambiguus var. salmonis tous nature and the widely spaced bunching habit Leymus salinus (Baker and Kennedy 1985). Elymus salina Salina wildrye spreads from seed and tillers. Rhi- zomes are more commonly produced under mesic con- Description ditions. On clay soils, rhizomes may be very short and inconspicuous. Salina wildrye is an erect, densely tufted perennial bunchgrass that occasionally produces short rhizomes. Plant Culture On some sites, particularly those with heavy clay soils, the plants form large tufts that give them the appear- Seeds of salina wildrye mature in mid to late July. ance of being nonrhizomatous. Plants are coarse Fruits are cleaned with an air screen machine. There stemmed with glabrous culms 1.1 to 3.3 ft (3.5 to 10 are about 130,000 seeds per pound (286,600 per kg) dm) tall. Leaves are glabrous to pubescent. Leaf blades (Link 1993). Seeds are estimated to remain viable for are flat or more commonly involute, 0.08 to 0.16 inch 2 to 3 years. Germination is enhanced by a 30-day (2 to 4 mm) wide, and often ciliate near the throat. moist prechill (Link 1993). Auricles are well developed and often clasping. Ligules Seedings of salina wildrye are often not successful are usually short, about 0.01 to 0.06 inch (0.2 to 1.5 due to low seed germination and poor seedling vigor. mm) long, squared apically, and ciliate along the Established stands, however, are persistent and margins. Spikes are erect, slender, 1.9 to 5.9 inches drought tolerant under a wide variety of conditions (5 to 15 cm) long, and 0.10 to 0.31 inch (2.5 to 8 mm) (Plummer and others 1968). The species has potential wide. Spikelets are usually solitary at each node and for increased use due to its wide range of adaptation, slightly overlapping. They are 0.35 to 0.79 inch (9 to 20 salt tolerance, drought resistance, and ability to stabi- mm) long and two to three or rarely six flowered. lize disturbed slopes with clay soils (Goodrich and Glumes are subequal, awl shaped, three nerved, smooth Neese 1986; Plummer 1977; Vallentine 1989). to roughened, sometimes broader at the base, and 0.05 to 0.39 inch (1.2 to 10 mm) long. There is often an extra Uses and Management glume at nodes producing only one spikelet. Lemmas are five-nerved (at least above), smooth to slightly Salina wildrye is moderately productive. Quality roughened on the surfaces, roughened on the awn and and palatability of the coarse forage is only fair when midnerve, and 0.28 to 0.41 inch (7 to 10.5 mm) long. green; it becomes unpalatable when cured (Vallentine Lemmas taper to a short awn that is 0.01 to 0.08 inch (0.2 to 2 mm) long. The species includes tetraploid (2n = 28) and octoploid (2n = 56) populations that do not differ morphologically. Plants flower in May to July (Arnow 1987, 1993; Cronquist and others 1977; Dittberner and Olsen 1983; Hitchcock 1950; Hitchcock and others 1969).

Ecological Relationships and Distribution Salina wildrye is distributed from California and southern Nevada to southwestern Wyoming. It is found from salt desert shrublands through desert shrub, sagebrush-grass, mountain mahogany, aspen, and conifer communities. It may be a common to dominant species in Gambel oak and pinyon-juniper communities (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969). Salina wildrye is drought resistant and adapted to dry Figure 29—Salina wildrye grows in mountainous ar- sites with loamy soils, but it also occurs on rocky to eas and on hillsides, alluvial fans, and canyons from sandy sites and on clay soils. It is moderately tolerant California to southwestern Wyoming (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 381 Chapter 18 Grasses

1989). Excessive grazing has resulted in declines of the species in northwestern Colorado (Baker and Kennedy 1985).

Varieties and Ecotypes The Upper Colorado Environmental Plant Center in Meeker, CO, is advancing ‘Accession 9043501’ from Colfax County, NM, toward release (UCEPC 2002). This accession has potential for revegetation of dry, moderately alkaline rangelands and other disturbed sites in the West. Its forage quality is moderate to fair for livestock and wildlife.

Elymus simplex Alkali Wildrye or Low-Creeping Wildrye ______Synonyms Elymus triticoides var. simplex Elymus simplex var. luxurians Leymus simplex Figure 30—Alkali wildrye spikes are elongate and Description slender with the spikelets generally solitary or sometimes two per node (Hitchcock 1950). Alkali wildrye is a waxy perennial that develops from extensively creeping rhizomes. Culms are 1.3 to 2.3 ft (4 to 7 dm) tall and the rhizomes can be up to 16.4 ft (5 m) long. Leaves are mostly basal. Leaf sheaths are hairless. The blades are usually involute to more or less flat, 0.04 to 0.12 inch (1 to 3 mm) wide, also confused with salina wildrye plants that produce rigid, and roughened or sparsely hairy on the upper rhizomes. Location is the best way to distinguish surface. Auricles are poorly developed and usually less between these two. Alkali wildrye is a grass of sandy than 0.04 inch (1 mm) long. Ligules are very short, bottomlands, whereas salina wildrye usually grows on about 0.02 inch (0.5 mm) long, squared at the apex, mountainsides and in heavy clay soils (Arnow 1987, and ciliate along the margins. Spikes are 2.4 to 6.3 1993; Cronquist and others 1977; Hitchcock 1950; inches (6 to 16 cm) long, erect, and usually rather Hitchcock and others 1969). slender (fig. 30). Spikelets are usually solitary, or sometimes two per node. When two per node, one is Ecological Relationships and Distribution sessile and the other is short pedicellate. Spikelets are 0.43 to 0.80 inch (11 to 20 mm) long, three to ten Alkali or low-creeping wildrye is found in moist, flowered, sometimes well spaced and not overlapping, sandy soils along riverbanks, and sometimes in drift- but usually slightly overlapping. Glumes are subequal, ing sands along the Green River in Daggett County, 0.22 to 0.63 inch (5.5 to 16 mm) long, awl shaped to only UT. It also occurs in Wyoming and Colorado (Arnow slightly broader at the base, hairless to roughened, 1987, 1993; Cronquist and others 1977). nearly nerveless, or one to three nerved. There is sometimes an extra glume at a one-spikelet node. Plant Culture, Uses, and Management Lemmas are 0.30 to 0.37 inch (7.5 to 9.5 mm) long, usually hairless to minutely pubescent and shiny Alkali wildrye has received little use in revegetation green or rarely with a purplish tinge. They are nerveless efforts. The species could be used on native sites to below and faintly five to seven nerved above. Glumes provide cover, forage, soil stabilization, and diversity. taper to a 0.10- to 0.31-inch (2.5- to 8-mm) awn. Plants flower from June to August. Varieties and Ecotypes Alkali wildrye is sometimes confused with small There are no releases. plants of creeping wildrye with solitary spikelets. It is

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Elymus triticoides is especially common in the Humboldt River drainage of Nevada (Jensen and others 2001). It is a long-lived, Creeping Wildrye or Beardless salt-tolerant, sodforming perennial that grows in dry Wildrye ______to moist places, valley bottoms, saline seeps and meadows, subirrigated sites including alkaline areas with Synonyms high water tables, and sites exposed to spring flooding. Elymus condensatus var. triticoides It occurs on saline meadows and salt desert shrub Leymus triticoides lands receiving 8 to 12 inches (20 to 30.5 cm) of annual Elymus simplex var. luxurians rainfall. The species is cross-pollinating and frequently Elymus acicularis does not set seed. Stands are generally not extensive; patches may represent single clones (fig. 31). Creeping Description wildrye often forms natural hybrids with Great Basin wildrye and salina wildrye (Arnow 1987, 1993; Creeping or beardless wildrye is a waxy blue or Cronquist and others 1977; Hitchcock 1950; Hitchcock sometimes green perennial that produces extensive and others 1969; Knapp and Wiesner 1978). and aggressive creeping rhizomes. Culms are 0.5 to 3.9 ft (1.5 to 12 dm) tall. Leaf sheaths are glabrous, Plant Culture roughened, or sometimes minutely hairy. Leaf blades are flat or more commonly involute, and measure Creeping wildrye seeds mature between midsum- 0.10 to 0.28 inch (2.5 to 7 mm) in width. They are mer and early fall (Sampson 1924). Fields may be coarse, stiff, roughened, and sometimes glabrous be- combined or swathed and windrowed. Seed retention low. Auricles are well developed and often clasping. at maturity is good, but lodging can interfere with Ligules are short, 0.01 to 0.28 inch (0.2 to 0.7 mm) long, harvest. Yields from seed fields average 100 to 200 squared, and irregularly lacerate and ciliate along the pounds per acre (112 to 224 kg per ha) (Smith and margins. Spikes are slender and about 1.4 to 3.5 inches Smith 1997). Stand life is about 5 years. Seeds are (3.5 to 9 cm) long, erect, loose, open to rather dense, cleaned with a fanning mill. There are about 175,000 and sometimes compound. Spikelets are paired or seeds per pound (385,000 per kg) (Smith and Smith sometimes solitary at the nodes, and 0.39 to 0.70 inch 1997). Dormancy results from an impermeable outer (10 to 18 mm) long. They are three to eight flowered seed coat. Germination is hastened by a 5-day prechill and greenish, brownish, or purplish. Glumes are at 1.5 ∞C (Wagner and Chapman 1970) with incuba- subequal, 0.16 to 0.43 inch (4 to 11 mm) long, narrow tion at 15/20 ∞C (Wagner and Chapman 1970) or 20/ and awl shaped, firm, one to three nerved, and mostly 30 ∞C (16 hrs/8 hrs) (Haferkamp and McSwain 1951). roughened. Lemmas are small, 0.24 to 0.39 inch (6 to Creeping wildrye is fall planted to provide overwinter 10 mm) long, often shiny and smooth, hairless, mi- exposure to cool, moist seedbed conditions. Seeds are nutely hairy, or sometimes minutely hairy only at the drilled or broadcast and covered. Slow germination, apices. They are faintly to prominently five to seven nerved, rounded on the back or keeled toward the tip, and awnless or with a short awn about 0.02 to 0.10 inch (0.5 to 2.5 mm) long. Plants flower from May through August. Considerable ecotypic variation occurs within the species. Creeping or beardless wildrye can be confused with western wheatgrass when there is only one spikelet per node. Very narrow glumes and a twisted rachilla that cause the florets to be out of their normal position are characteristic of creeping wildrye (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969), and although of differing growth habits, creeping wildrye and Great Basin wildrye share similar genomes and hybridize in nature (Dewey 1984). Creeping wildrye also hybridizes with salina wildrye (Arnow 1987).

Ecological Relationships and Distribution Figure 31—Patches of creeping wildrye often represent individual clones (photo courtesy of Tom Creeping wildrye is distributed from Washington to Jones, USDA ARS, Logan, UT). Montana, and south from Baja California to Texas. It

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 383 Chapter 18 Grasses low seedling vigor, and resulting poor establishment five to seven faint nerves. Awns, when present, are up have limited use of this species. Although young stands to 0.08 inch (2 mm) long. Plants flower from late May are not competitive, established stands are highly through July (Arnow 1987, 1993; Cronquist and oth- persistent. ers 1977; Hitchcock 1950; Hitchcock and others 1969). The species is allohexaploid with 2n = 6n = 42 (Sleper Uses and Management and West 1996). Creeping wildrye is seeded to provide soil stabiliza- Ecological Relationships and Distribution tion on highly erodible sites and can be used for reseeding disturbances along waterways and on mined Tall fescue, native to Europe, is seeded as hay, lands, roadways, and disturbed rangelands. It is one of pasture, or turf grass in the Western Hemisphere. The the few species that can be seeded to provide forage on species contains considerable variability, and many wet, alkaline sites in low-precipitation areas, and it is cultivars are available. Leaves of the forage varieties useful in saline areas and areas that are flooded in are generally soft and lax compared to those of the turf spring. It can be used in native species restoration varieties (Jensen and others 2001). Tall fescue is efforts in salt desert shrub and pinyon-juniper com- cultivated from Alaska and southern Canada through- munities. Native stands provide cover and forage for out much of the United States (fig. 32). It is also wildlife and are sometimes cut for hay (Jensen and present in South America. The species has escaped others 2001). and become widely established in areas where it has been seeded. Tall fescue can now be found in seeded Varieties and Ecotypes fields, moderately moist areas along irrigation ditches and roadsides, and in fallow fields and seeded moun- ‘Rio’ was released by the Lockford Plant Material tain meadows. Its best growth is on deep, moderately Center in 1991. It was selected for its vigorous spread- fertile, fine silty to clay soils, but it will grow well on a ing habit. ‘Shoshone’, released in 1980 by the Bridger wide range of soil textures when moisture is adequate Plant Materials Center, is a productive and rhizoma- (Wasser 1982). It can also be seeded on moderately tous cultivar. saline and alkaline soils (Jensen and others 2001). Growth is good in areas that are irrigated, subirrigated, Festuca arundinacea or that receive at least 18 inches (460 mm) of annual precipitation (Jensen and others 2001; Wasser 1982). Tall Fescue, Reed Fescue, or Alta Tall fescue is a relatively drought- and heat-tolerant Fescue ______Synonyms Bromus arundinaceus Festuca elatior var. arundinacea

Description Tall fescue is a stout, often strongly tufted perennial with an extensive coarse, fibrous root system. Rhi- zomes are generally short and vary considerably in their sodforming ability. Culms are 1.6 to 6.6 ft (0.5 to 2 m) tall and ascending or erect with smooth or roughened leaf sheaths. Leaf blades are stiff, flat or somewhat involute, 0.12 to 0.47 inch (3 to 12 mm) wide, and roughened above. Auricles are prominent and usually ciliate. Ligules are about 0.01 inch (0.2 mm) long. Panicles are narrow, 5.9 to 13.8 inches (15 to 35 cm) long, and often somewhat nodding. Spikelets measure 0.4 to 0.6 inch (10 to 15 mm) in length and are five to nine flowered. Glumes are lance shaped. The Figure 32—Reed fescue grows on moist soils and first is 0.16 to 0.24 inch (4 to 6 mm) long and one to tolerates saline and alkaline conditions. Grown for three nerved; the second is 0.16 to 0.28 inch (4 to 7 mm) pasture and hay production, it has escaped and long and three to five nerved with translucent mar- become widely distributed in North America (RMRS gins. Lemmas are 0.3 to 0.4 inch (7 to 10 mm) long with photo).

384 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses pasture grass, but it does not survive long dry periods. They are particularly useful on saline and heavy This species tolerates cold, winter flooding, high water alkaline soils where few other grasses can be seeded tables, and poorly drained soils, and it is moderately (Arnow 1987). The species is persistent, palatable, shade tolerant. Plants are resistant to burning when moderately to highly productive, and compatible with dormant, but the species does not commonly occur in a variety of management programs (Hanson 1979). areas susceptible to wildfire. Tall fescue can be grown with selected legumes if An endophytic fungus, Acremonium coenophialum, managed carefully (Wasser 1982). When grown alone, frequently infects tall fescue (Sleper and West 1996). it requires added nitrogen and irrigation to maximize When infected forage is consumed in large quantities, productivity (Thornburg 1982). Tall fescue is tolerant susceptible livestock may develop lameness and some- of periodic or continuous grazing and can be grazed or times gangrene (Kingsbury 1964). Endophyte-free mown to a 2- to 4-inch (5 to 10 cm) stubble height cultivars should be used for forage production, but (Plummer and others 1968; Wasser 1982). Grazing these often require more fertile soils or fertilizer appli- should be applied evenly to maintain a uniform stand. cations. Tall fescue is also an invader of lawns (Arnow Frequent grazing improves palatability, induces new 1987, 1993; Cronquist and others 1977; Hitchcock tiller production, and maintains a dense sod (Wasser 1950; Hitchcock and others 1969). 1982). Stands are more resistant to trampling than most pasture grasses. Plant Culture Tall fescue provides a long grazing season. It receives greatest use in spring and early summer when Seeds are harvested by direct combining or by swath- leaves are young. Its productivity and palatability are ing and combining. Some seed loss normally results greatly reduced in late summer when the leaves be- from shattering. Seeds are cleaned using an air-screen come coarse (Hassell and others 1983). Summer and machine. There are 181,000 to 230,000 seeds per fall growth retain digestibility and crude protein con- pound (399,000 to 507,100 per kg). Minimum purity tent better than many cool-season grasses (Matches for seed purchases is 98 percent, and germination is 85 1979). Leaves remain green and provide good-quality percent (Wasser 1982). Seeds are drilled or broad- fall and winter forage for livestock. Tall fescue pro- casted seeded and covered to 0.25 to 0.5 inch (0.6 to 1.3 vides excellent quality hay (Jensen and others 2001). cm) deep depending on soil conditions. Germination Tall fescue increases over time when planted with and initial seedling establishment and growth are other pasture grasses due to its lower palatability moderate to good (Hassell and others 1983; Plummer (Jensen and others 2001). Cutworms, slugs, and grass- and others 1968). Seedlings usually become estab- hoppers can reduce yields. lished after one growing season (Jensen and others Tall fescue has been seeded to provide forage in 2001). Plants are persistent, good seed producers, and mountain meadows and in openings in aspen and spread well from seed. Seed production is centered in conifer forests (Vallentine 1971). It is used by elk and the Northwest, and yields of 2,510 pounds per acre deer, but its palatability to big game is generally low. (2,240 kg per ha) are obtained in western Oregon Seeds are consumed by songbirds, and the seeds and (Sleper and West 1996). foliage are used by small mammals. Sodforming culti- Irrigated tall fescue fields are planted in late sum- vars are used to stabilize soils on road cuts and fills, mer. Weed infestations must be controlled until plants and disturbances in riparian areas and recreational are established. When legumes are added to improve areas, and to provide cover crops. Fertilizer applica- forage palatability and quality, seeding may be done tions may be required when using the species on in early spring (Jensen and others 2001). The legumes infertile soils (Vallentine 1967). are often planted in alternate rows with the grass. Ladino clover, alfalfa, white clover, and birdsfoot tre- Varieties and Ecotypes foil are used for this purpose (Sleper and West 1996; Wasser 1982). Annual nitrogen applications are re- ‘Alta’ and ‘Fawn’ were developed from European quired when tall fescue is seeded without legumes. material (Alderson and Sharp 1994) and are adapted Wildland sites in mountainous areas are seeded in to valley bottoms where they are seeded as pasture late fall or early summer. The resulting stands are grasses and for the renovation of saltgrass sites. Both competitive and persistent when established, but they varieties are also used for soil stabilization and forage may be somewhat slow to develop. New seedings production in semiwet meadows of upper elevation should not be grazed until plants are well established areas and in openings in aspen and conifer forests. They require at least 16 inches (410 mm) of annual Uses and Management precipitation. Both green up early in spring. They are compatible with other species and can be included in Tall fescue seedings provide pasture or hay in irri- mixtures. gated fields in valleys of the Intermountain region.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 385 Chapter 18 Grasses

Several tall fescue varieties have been developed for Panicles are 1.9 to 5.9 inches (5 to 15 cm) long, loosely hay and pasture forage production. These include compressed, and extending mostly from one side of the ‘Barcel’, ‘Barvetia’, ‘Cajun’, ‘Courtenay’, ‘Dovey’, rachis. Spikelets are 0.31 to 0.51 inch (8 to 13 mm) long ‘Festorina’, ‘Fuego’, ‘Georgia No. 5’, ‘Goar’, ‘Kentucky and four to seven flowered with visible joints on the 31’, ‘Martin’, ‘Maximize’, ‘Mozark’, ‘Penngrazer’, axis of the spikelet. Glumes are narrowly lanceolate ‘Phyter’, ‘Southern Cross’, ‘Stargrazer’, and ‘Stef’ and taper to a pointed or rounded apex. The first glume (Alderson and Sharp 1994). None of these varieties is 0.10 to 0.18 inch (2.5 to 4.5 mm) long, the second is have proven adapted to mountain rangeland condi- 0.16 to 0.22 inch (4 to 5.5 mm) long. Lemmas are 0.20 tions. There are a number of other varieties that have to 0.28 inch (5 to 7 mm) long, rounded, and glabrous to been bred for use as turf grasses. roughened distally. Awns are 0.08 to 0.24 inch (2 to 6 mm) long. Plants flower from May to August (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock Festuca idahoensis 1950; Hitchcock and others 1969). Idaho fescue is very Idaho Fescue or closely related to Festuca ovina. Some authors regard Bluebunch Fescue ______it as the native, low-elevation phase of that species and apply the name F. ovina var. ingrata. Synonyms Ecological Relationships and Distribution Festuca ovina var. ingrata Festuca ingrata Idaho fescue ranges from Southwestern Canada Festuca occidentalis var. ingrata south to northern California and east from central Festuca ovina var. oregona Montana to northern New Mexico (Tisdale 1959). It Festuca ingrata var. nudata occurs at elevations ranging from 800 to 12,000 ft (244 Festuca ovina var. nudata to 3,659 m) on sites receiving 15 to 30 inches (380 to 760 mm) of annual precipitation (Daubenmire 1966). Description On north slopes and cool areas receiving added moisture it may grow where annual precipitation is as low Idaho fescue is one of the most common bunch- as 10 inches (254 mm). Idaho fescue occurs over a wide grasses in the West. It is a densely tufted environmental gradient and is associated with a range nonrhizomatous, long-lived perennial bunchgrass with of soil conditions and plant communities. It grows on hairless to roughened culms that grow 1 to 3.3 ft (3 to dry, well-drained, fertile, moderately deep soils rang- 10 dm) tall (fig. 33). Leaves are numerous, fine, and ing from clayey or silty to sandy and gravelly loams mostly basal, forming a tuft 6 to 12 inches (15 to 30 cm) that are slightly acidic to slightly alkaline (USDA tall that is usually more than half the length of the Forest Service 1937; Wasser 1982; Vallentine 1971). It culms. Leaf blades are firm, involute, less than 0.04 also occurs on shallow, rocky soils and on weakly inch (1 mm) wide, and without auricles. Ligules are saline soils where its productivity is lower. Plants are mostly 0.01 to 0.02 inch (0.3 to 0.6 mm) long, finely cold tolerant and moderately drought tolerant, but ciliate, and longer on the sides than at the center. they are not tolerant of high water tables or flooding. Idaho fescue is a common or codominant species in grasslands, sagebrush-grass, mountain mahogany, oakbrush, mountain brush, pinyon-juniper, antelope bitterbrush, mountain meadow, conifer, riparian, and alpine communities (Zouhar 2000). It occurs on meadows, plains, foothills, ridgetops, mountain brush communities, and open conifer forests. The species occurs in open areas, but it is also moderately shade tolerant. Associated species are big sagebrush, Sandberg bluegrass, bluebunch wheatgrass, and many native forbs. In lower elevation and drier communities, the distribution of Idaho fescue is restricted to areas with better availability of water (Johnson 1994). Many Idaho fescue sites have been invaded by cheatgrass (Wasser 1982). Seedlings may emerge in fall and develop during Figure 33—Idaho fescue is one of the most impor- favorable periods in winter and spring to the point that tant cool-season bunchgrasses in the West (RMRS the plants can become dormant and survive summer photo).

386 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses drought (Daubenmire 1975b). Idaho fescue begins the hard-dough stage. Yields are about 300 pounds per growth in March or April, and seeds mature in mid to acre (336 kg per ha) from irrigated fields and 100 late summer depending on location and altitude. Plants pounds per acre (112 kg per ha) from nonirrigated become dormant during summer, but resume growth fields. A debearder will remove awns; seeds are cleaned in fall if adequate moisture is received (Wasser 1982). using a fanning mill (Smith and Smith 1997). There Reproduction is by seed and tillers (Stubbendieck are about 425,000 to 460,000 seeds per pound (937,000 1992). Plants produce shallow, fibrous root systems. to 1,014,100 per kg) (Ensign and others 1984). Recom- Goodwin and others (1996) found that about 40 per- mended quality standards are 90 percent purity and cent of the root biomass occurs in the upper 4 inches 80 percent germination (Wasser 1982). Seeds undergo (10 cm) of soil. Roots are associated with vesicular- afterripening over a period of at least 6 months arbuscular mycorrhizae (Goodwin 1992; Ho 1987). (Goodwin and others 1996), thus germination is de- Idaho fescue can be damaged by wildfires at any layed until seeds have gone through at least one season (Smith and Busby 1981; Wright and others winter. Idaho fescue can be germinated by incubating 1979). Cool fires burning through stands with little the seeds at 50/68 ∞F or 68/86 ∞F (10/20 ∞C or 20/30 ∞C) litter buildup may cause minimal damage. However, (16/8 hrs) (Haferkamp and McSwain 1951). Exposure the fine leaves and accumulation of litter can result in to light improves germination (Haferkamp and prolonged high-temperature burning that may kill or McSwain 1951). damage the root crown (Agee 1996), with the extent of Idaho fescue is fall seeded on drier sites by broad- damage dependent upon fire conditions, fire history of casting or drilling on a firm seedbed. Seed should be the site, extent of grazing, and other factors (Bunting planted about 0.5 inch (1.3 cm) deep. In irrigated seed 1984; Wright and others 1979). Rate of postfire recov- fields or on low-elevation wildland sites that receive ery varies considerably among burns and depends on reliable moisture it can be spring seeded. Dry sites are the degree of plant damage and environmental factors fall seeded. At high elevations it can be spring or early (Beardall and Sylvester 1976; Harniss and Murray summer seeded as soon as the site is open. 1973), with response to season of burn variable. Sur- In seed fields, drilling into stubble or planting in viving plants recover through tillering if soil moisture alternate rows with annual grains may establish seed permits. Both protein content and digestibility of Idaho fields. Fallowing also reduces weed populations. Row fescue increase temporarily following burning (Beardall spacings of 30 to 36 inches (76 to 92 cm) are recom- and Sylvester 1976; Singer and Harter 1996). Fires mended for seed fields (Wasser 1982). Expected stand near the margins of the species range are damaging at life is about 5 years (Smith and Smith 1997). any season (Blaisdell 1953; Wright and others 1979); Vigor of seedlings is poor to fair (Shaw and Cooper fires in dry environments tend to reduce productivity 1973; Vallentine 1971), and they are not competitive for longer periods because the shallow root systems with seedlings of more aggressive species (Hafenrichter are heavily damaged. Seedling establishment is re- and others 1968). Stands require 2 to 3 years to duced following burns, likely due to reduction of the establish (Hafenrichter and others 1968; Plummer seed bank when temperatures exceed 257 ∞F (125 ∞C) and others 1968), and weeds must be controlled in seed for more than 5 minutes (Warg 1938). It is not clear fields. Grazing should be restricted until stands are whether early- or late-season burning best protects established (Ensign and others 1984); grazing pres- Idaho fescue plants, but Wright (1974) recommended sure must be carefully controlled to maintain stands. burning when species of concern are dormant and not Resting during the season or grazing on a rotational stressed by drought. Bunting and others (1998) found basis may be required (Hormay and Talbot 1961). that deferring grazing for at least 1 year following Seedlings are susceptible to attacks by damping off burning improved recovery of Idaho fescue. and other seedling diseases, grasshoppers, and rodents (Wasser 1982). Competitive ability increases Plant Culture considerably as Idaho fescue stands mature. When seeded on erosive sites, addition of mulch may be Variable and often low seed production and viability required to stabilize the surface soil until plants be- in wildland stands has limited the use of Idaho fescue come established (Hafenrichter and others 1968). in wildland seedings (Ensign and others 1984; Wasser 1982). Overcoming this problem has been a major Uses and Management emphasis in plant selection work with the species; two cultivars are now available. Lodging is not nor- Idaho fescue is very palatable and heavily used by mally a problem in seed fields. Harvesting is accom- livestock and wildlife. It is a particularly valuable plished by swathing before seed shattering occurs in species at mid to higher elevations in sagebrush early to midsummer. Seed can be combined when in grasslands, open conifer forests, and alpine areas.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 387 Chapter 18 Grasses

Greatest use is received in spring when the forage is Festuca ovina succulent and palatable, but use, particularly by sheep, declines later in the season as the forage becomes Sheep Fescue or coarse and dry. Animals will continue to use it if other Alpine Fescue ______forage is not available. Plants cure well and provide good fall and winter forage where accessible (Shaw Synonyms and Cooper 1973). The species withstands some use, Festuca brevifolia but stands are weakened or lost with excessive grazing Festuca minutiflora and trampling, particularly in early spring. Stands Festuca saximontana maintained in healthy condition will reseed themselves. Deer use Idaho fescue in spring (Johnson 1994), Description and elk use it throughout the year (Dragt and Havstad 1987; Kufeld 1973; Shaw and Cooper 1973). Sheep fescue is a short, matforming perennial with Idaho fescue can be planted alone or in mixtures dense tufts of basal leaves. Culms are 0.3 to 1.6 ft (0.8 with other native grasses to provide diversity. Al- to 5 dm) tall. The tufts are 2.0 to 5.9 inches (5 to 15 cm) though not deep rooted, it produces an extensive, tall and generally about half the height of the culms. fibrous root system that provides soil stabilization and Sheaths are glabrous to minutely hairy. Leaf blades ground cover on disturbances resulting from logging, are stiff, involute, and without auricles. Ligules are road construction, mining, and recreation (Ensign and scarcely 0.02 inch (0.4 mm) long, finely ciliate, and others 1984; Hafenrichter and others 1968). It can be longer on the sides than at the center. Panicles are 0.6 seeded in areas where tree seedlings have been planted to 3.95 inches (1.5 to 10 cm) long, mostly strongly if the seeding rate is low to moderate (Ensign and compressed, but somewhat open in F. o. var. ovina. others 1984). Mature stands withstand or slow the Spikelets are 0.16 to 0.35 inch (4 to 9 mm) long and two spread of invasive species (Borman and others 1990; to five flowered. Glumes are narrowly to broadly lan- Hafenrichter and others 1968; Lilley and Benson 1979). ceolate. The first glume is 0.08 to 0.14 inch (2 to 3.5 mm) long and one nerved. The second is 0.12 to 0.18 Varieties and Ecotypes inch (3 to 4.5 mm) long and three nerved. Lemmas are 0.14 to 0.24 inch (3.5 to 6 mm) long, glabrous or The varieties ‘Joseph’ and ‘Nezpurs’ were developed minutely hairy toward the tip. Awns are 0.04 to 0.20 from material collected in the Northwestern United inch (1 to 5 mm) long. Sheep fescue flowers in July States and Southwestern Canada (Ensign and others through August and disperses its seed from August to 1984). They are adapted to elevations ranging from October. Reproduction is solely by seed due to the 1,000 to 9,000 ft (305 to 2,745 m) and communities absence of rhizomes (Arnow 1987, 1993; Cronquist ranging from sagebrush grass to subalpine that re- and others 1977; Hitchcock 1950; Hitchcock and oth- ceive 14 to 30 inches (355 to 760 mm) of annual ers 1969). precipitation. Both varieties are cold and drought hardy and moderately shade tolerant. They are best Ecological Relationships and Distribution adapted to silty to sandy loam soils that are slightly alkaline to slightly acidic. They do especially well in Sheep fescue is a widespread, polymorphic ponderosa pine and mountain big sagebrush commu- circumboreal species with many varieties, some of nities on gravelly to well-drained loamy soils. Joseph which have been treated on the specific level. In North and Nespurs are seeded on summer ranges to provide America, sheep fescue is distributed southward from forage for livestock and wildlife and soil stabilization the arctic region through mountainous areas to Cali- on road cuts and fills, skid trails, cutover forest areas, fornia, and east to northern New Mexico, Kansas, and and recreational sites. These varieties perform well as North Carolina. The species also occurs in Mexico and components of seeding mixes because they are not South America. Plants of the two native varieties are overly competitive. small and occur in rocky areas and on shallow, grav- Joseph was selected for its greater leaf and culm elly soils, and dry exposed sites on slopes and ridges. height (28 to 31 inches [72 to 80 cm]), basal leaf growth, They require annual precipitation of 8 to 14 inches (20 and good germination (Ensign and others 1984). to 36 cm) or more (Arnow 1987), and may be abundant Nezpurs was selected for its shorter stature, 20 to 28 in communities from sagebrush to pinyon-juniper, inches (50 to 70 cm), large seed size, productivity, mountain brush, aspen, conifer, and mountain grass- basal leaf growth, and olive-green color. Both varieties land communities. F. o. var rydbergii is distributed have been seeded in the Intermountain West, especially from foothill to subalpine areas and F. o. var. brevifolia in Idaho, Wyoming, eastern Oregon, and western occurs in subalpine and alpine habitats (Cronquist Colorado. Nezpurs has received more use than Joseph. and others 1977).

388 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

The three varieties commonly seeded are all Euro- of annual precipitation (Wasser 1982) to stabilize soils pean in origin: sheep fescue, hard sheep fescue) and revegetate disturbances. It is used for roadway (fig. 34), and sulcate sheep fescue (fig. 35). Sheep plantings, airports, and other sites where a competi- fescue is distinguished by its open and spreading tive, low-growing ground cover is required. Although panicle. Hard sheep fescue has leaf blades that are seedlings are slow to establish, plants are long lived smoother, wider, and firmer than those of sulcate and become competitive as they develop through the sheep fescue. growth of abundant fibrous roots. Forage production is Sheep fescue is a long-lived, drought-hardy variety. low, but of fair to good quality (Hassell and others It grows on coarse, dry, infertile soils on sites receiving 1983). Sheep fescue is adapted to shallow, moderately 10 to 18 inches (25 to 46 cm) of annual precipitation to well-drained soils with pH 5.5 to 7.5. Plants are heat (Jensen and others 2001). Sheep fescue is cold tolerant tolerant. and provides stability to surface soils. It is somewhat tolerant of finer soils and shading. Hard sheep fescue Plant Culture is seeded on sites receiving 15 inches (38 cm) or more Sheep fescue seeds ripen in early fall (Sampson 1924). They can be combined directly or first windrowed and dried for 5 to 7 days (Smith and Smith 1997; Wheeler and Hill 1957). Lodging is generally not a problem, but seeds begin shattering after reaching the hard-dough stage (Smith and Smith 1997). Seed is cleaned using a debearder and fanning mill. An indent cylinder may be used to increase purity. Seeds are not generally harvested during the establishment year. Stand life is 4 to 5 years. There are about 500,000 seeds per pound (1,100,000 per kg) (Smith and Smith 1997). Seed fields can be planted in fall or spring (Jensen and others 2001; Plummer and others 1968). Seeds are easily handled. They should be planted about 0.25 to 0.5 inch (0.6 to 1.5 cm) deep (Horton 1989; Plummer 1943) on a firm seedbed. Rows should be 12 or 24 inches (30 to 60 cm) apart in irrigated fields and 36 inches (90 cm) apart on drylands (Smith and Smith 1997). Wildland sites are drill seeded or broadcast. Dry sites are best seeded in fall. More mesic areas can Figure 34—Hard sheep fescue plants are upright, be spring seeded. Seedlings are not highly vigorous, leafy, and slightly taller than those of sulcate sheep and establishment occurs over a 2- to 3-year period as fescue (RMRS photo). plants develop extensive fibrous root systems. On wildland sites, sulcate sheep fescue is more compatible with other species than hard fescue and can be included in mixtures. Plants are not tolerant of highly saline or alkaline soils (Jensen and others 2001); they are useful on dry, sandy, or rocky sites (USDA Forest Service 1937) where other components of the mixture may not establish well. Mature plants are persistent due to development of an extensive fibrous root system. Plantings reduce the spread of weeds and are tolerant of grazing if properly managed. Excessive grazing can damage the stands; recovery is slow (Jensen and others 2001).

Uses and Management Sheep fescue has been seeded successfully throughout the Intermountain West in mountain big sagebrush and mountain brush communities and in open- Figure 35—Sulcate sheep fescue grows as dis- ings in aspen and subalpine communities that receive tinct fine-leaved clumps (RMRS photo). 14 inches (355 mm) or more of annual precipitation. Its

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 389 Chapter 18 Grasses seedlings are slow to establish, but tenacious. The (fig. 36). Nodes of the culms are often finely hairy. extensive root system of mature plants makes them Leaves are mostly basal with hairless to minutely competitive and fairly resistant to drought and tram- roughened sheaths, and a few shaggy hairs at the pling. Sheep fescue greens up early in spring and throat. Leaf blades are short and curly, flat to folded, receives some use by livestock, particularly sheep. 0.06 to 0.12 inch (1.5 to 3 mm) wide, 0.8 to 2 inches (2 Grazing should be regulated to leave a 2- to 4-inch (5 to 5 cm) long, and involute toward the tip. They are to 10 cm) stubble (Jensen and others 2001). rigid, hairless to roughened, often waxy, and some- Sheep fescue is used for postfire seedings, mined times covered with small stiff hairs. Ligules are 0.04 to land reseedings, recreation areas, ditchbanks, and 0.10 inch (1 to 2.5 mm) long with lacerate, long-ciliate other disturbances in mountainous areas to reduce margins. Spikes are 0.08 to 0.28 inch (2 to 7 mm) in soil erosion. Baron (1962) found conifer seedling sur- length with long, shaggy hairs at the base of the vival was not decreased on areas planted with sheep spikelet clusters. Glumes of the lateral spikelets are fescue relative to controls. Sheep fescue is used in lawn subequal, roughened, and 0.20 to 0.33 inch (5 to 8.5 seed mixtures and is particularly useful on infertile or mm) long with an irregularly lacerated apex. The first sandy soils. glume is asymmetrical with the midnerve displaced to one side; awn length is 0.12 to 0.22 inch (3 to 5.5 mm). Varieties and Ecotypes The second glume is awnless, or very short awned. Glumes of the central spikelet are subequal and 0.16 ‘Covar’ sheep fescue originated near Konya, Turkey to 0.26 inch (4 to 6.6 mm) long with stiff hairs. The (Alderson and Sharp 1994). A fine-leaved fescue, it nerves in the glumes extend into an irregular awn that was selected for erosion control and revegetation of measures 0.06 to 0.28 inch (1.5 to 7 mm) in length. disturbed sites and depleted communities. It is adapted Lemmas of the lateral and central spikelets are 0.18 to to shallow, gravelly, and well-drained soils that re- 0.35 inch (4.5 to 9 mm) long, lanceolate, and blunt ceive at least 8 inches (203 mm) of annual precipita- tipped. They are hairless basally, roughened apically, tion (Wasser 1982). Covar has been seeded success- and three nerved. The single lemma of the central fully in mountain big sagebrush, upper mountain spikelet is narrowly lanceolate and bears a dorsal awn brush, ponderosa pine forests, and aspen openings. It from below the notched apex (Arnow 1987, 1993; requires fall seeding, and it is slow to establish. Ma- Cronquist and others 1977; Hitchcock 1950; Hitchcock ture plants are persistent, competitive, winter hardy, and others 1969). Galleta is sometimes referred to in and drought tolerant. The variety is competitive and older literature as black grama. This common name is will spread to dominate a community. misleading, giving the impression that it belongs to ‘Durar’ hard fescue, a larger plant of European the genus Bouteloua. Galleta flowers from May to origin (Alderson and Sharp 1994), is adapted to well- August. Chromosome number is 2n = 2x = 36 or 38 drained, basic to moderately acidic soils of pinyon- (Arnow 1987). juniper, mountain brush, ponderosa pine, aspen, and conifer forests receiving 14 to 28 inches (300 to 700 mm) of annual precipitation. The variety was developed for soil stabilization and forage production. Durar is somewhat slow to establish, but mature stands spread aggressively and outcompete most native and some introduced grasses, forbs, and shrubs.

Hilaria jamesii Galleta or Curly Grass ______Synonyms Hilaria sericea

Description Galleta is a strongly rhizomatous or stoloniferous perennial warm-season grass with a dense, fibrous root system. Rooting depth is about 12 inches (30 cm). Figure 36—A strongly rhizomatous or stoloniferous Culms are 0.5 to 2.1 ft (1.5 to 6.5 dm) tall and grow species, galleta stems grow along the ground before horizontally along the ground before turning upward bending upward (RMRS photo).

390 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Ecological Relationships and Distribution designed to plant chaffy seed. Seed may be broadcast, but it must be covered. Light irrigation helps to increase Galleta is the most northerly distributed species of germination and establishment in seed fields. its genus (Heizer and Hassell 1985). Its range extends Wildland stands may be established using water from southeastern California and southern Nevada catchment techniques such as deep furrow drilling to across Arizona and southern Utah to Wyoming, New aid stand establishment on dry sites. Rate of seedling Mexico, Kansas, Texas, and the Oklahoma Panhandle development is slow to moderate and stand establish- (Heizer and Hassell 1985). It grows from dry desert ment may require 2 years or more. Seedlings are scrublands of lower valleys to mesas, plains, and drought and cold tolerant, but they are susceptible to pinyon-juniper woodlands of foothills. Galleta often attacks by rodents and grasshoppers (Wasser 1982). grows in association with blue grama in sagebrush Galleta is adapted to fine, sometimes saline soils that areas, often as the dominant species. It occurs from the limit seedling establishment of many species. It can be valleys to the lower ponderosa pine zone at elevations seeded in mixtures with other species adapted to such from 4,000 to 8,000 ft (1,220 to 2,439 m) (USDA Forest sites. Grazing should be avoided during one or more Service 1937; Wasser 1982). It is particularly common growing seasons to permit stand establishment (Merkel and frequently dominant on the plains of New Mexico. and Herbel 1973). It grows with shadscale on salt desert shrub sites in the Great Basin and on sandy and rocky sites in the Uses and Management Colorado Plateau. It also occurs in creosote bush and desert shrub communities in the Southwest. Galleta produces considerable nutritious forage and Galleta grows on sites receiving 6 to 18 inches (150 to withstands heavy grazing and drought. It is palatable 460 mm) of annual precipitation with soils ranging in pure stands, but receives less use when mixed with from coarse to fine and pH of 6.6 to 8.4 (Bridges 1942; more palatable grama grasses. When grazed heavily, Horton 1989; USDA Natural Resources Conservation such mixtures gradually shift toward a dominance of Service 2003; Stefferud 1948). Plants have a low re- galleta. Galleta provides good, moderately palatable quirement for soil fertility (USDA Natural Resources forage for livestock and wild ungulates during the Conservation Services 2003); tolerance of salinity and summer growing season. Regrowth is good following alkalinity varies among populations. Plants are summer rainfall, but regrowth following fall rains or drought and cold tolerant, but not shade tolerant. frost is more limited. Dormant plants are dry, tough, Galleta is moderately fire tolerant and usually recov- and unpalatable (Heizer and Hassell 1985; Vallentine ers by the second year following burning (Arnold and 1971; West 1972). Plants receive little use in late fall others 1964). or winter. Although galleta is resistant to grazing, excessive use results in reduced forage and rhizome Plant Culture production (Heizer and Hassell 1985). Galleta is seeded to provide forage, soil stabilization, Galleta initiates growth in spring if adequate mois- and watershed protection (Wasser 1982). Spread from ture is received (Wasser 1982). If summer rains occur, seed is slow, but the woody rootstocks that some- it may mature two seed crops. The first crop can be times grow to 6 or 7 ft (1.8 to 2.1 m) in length permit harvested in June and the second in August to Octo- vegetative spread. They also protect plants from ber. Many florets in wildland stands do not produce trampling damage and hold soils in erodible areas. seed, thus costs of wildland collections are high. The Plants are bunchy and do not form a continuous sod. release, ‘Viva’, was selected for its high percentage of Galleta is extremely drought resistant and provides a seed-bearing florets (Heizer and Hassell 1985). Native persistent vegetative cover in adapted areas. and domestic seed fields can be harvested by combining (Wasser 1982). Seed is cleaned by hammermilling Varieties and Ecotypes and fanning. Recommended purity is 20 to 40 percent and germination 80 to 98 percent for seed sales. There Viva was collected from a population near New Kirk, are about 170,000 seeds per pound (374,780 per kg). NM, on a site at 5,740 ft (1,750 m) elevation that Seeds do not require prechilling. They can be incu- receives an average of 9 inches (230 mm) of annual bated at 68/86 ∞F (20/30∞C) (16/8 hrs) to test germina- precipitation (Alderson and Sharp 1994). It exhibits tion (Haferkamp and McSwain 1951). Exposure to good seedling vigor and drought and cold tolerance. light is required. Seeds are nondormant and will Viva provides forage for livestock and big game. It is germinate under a range of constant and alternating also used to revegetate mined lands and other distur- temperatures (Knipe 1967). bances. It is seeded in extreme southern Utah and Seed should be planted during the most reliable period Colorado, northern Arizona, and northwestern New of rainfall, usually early summer (Link 1993). Seed is Mexico. It has done well when seeded with other planted about 0.5 to 1 inch (1.3 to 2.5 cm) deep using drills warm-season grasses on the Great Plains from Mexico

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 391 Chapter 18 Grasses to the Canadian border. It has not established or to Texas, California, and Washington. It is one of the persisted well in the Great Basin. most common and widely distributed of the western grasses, extending over a wide range of altitudes and growing on a variety of dry to moist clay loam to sandy Koeleria macrantha or sometimes rocky soils with pH 6.5 to 8.0 (Hassell Prairie Junegrass ______and others 1983). Junegrass is an important mid- to late-seral forage species of the sagebrush, mixed grass, Synonyms and open timber types (Coupland 1950). It requires 16 Koeleria cristata to 21 inches (41 to 53 cm) of annual precipitation, and Poa cristata is usually found at elevations of 5,000 to 8,000 ft (1,524 Aira cristata to 2,438 m) (Parker 1975). The species does not yield a Koeleria gracilis great deal of forage, as it is low growing with abundant Koeleria nitida but short and mostly basal leaves. Prairie junegrass begins growth from seed or established plants very Description early in the season (Simonin 2000). Seed is dispersed from July to September. The generally low seed viabil- Prairie junegrass was originally and erroneously ity is counterbalanced by abundant production (Arnow named and described as K. cristata, based on a specimen of Poa cristata. The oldest valid name is believed to be K. macrantha; however, Cronquist and others (1977) prefer to use the oldest New World name, K. nitida. Junegrass can be distinguished from Poa fendleriana and Trisetum wolfii by its hairy panicle axis and generally smaller spikelets. It is often found in association with these grasses. Prairie junegrass is a cespitose perennial bunchgrass with culms 0.6 to 2.2 ft (2 to 6.5 dm) tall. Most leaves are basal, but a few small leaves are produced on the culms. Sheaths are stiff haired or rarely hairless. Leaf blades are usually folded or involute, 0.04 to 0.10 inch (1 to 2.5 mm) wide, and stiff haired to hairless. Ligules are 0.02 to 0.08 inch (0.5 to 2 mm) long, irregularly lacerate to subentire, minutely ciliate, and sometimes sparsely short hairy. Panicles are 1.0 to 5.9 inches (2.5 to 15 cm) long, contracted, and spikelike with short, appressed branches (fig. 37). The spike axis is densely short hairy. Spikelets are 0.16 to 0.20 inch (4 to 5 mm) long and two to four flowered. Spikelet axes are silky haired. Glumes are subequal and roughened on the keel and sometimes throughout. The first glume is 0.11 to 0.20 inch (2.8 to 5 mm) long, lanceolate to narrowly lanceolate, and one nerved. The second glume is 0.13 to 0.22 inch (3.2 to 5.5 mm) long, lanceolate, and one to three nerved with the lateral nerves barely visible. Lemmas are about as long as the second glume, lanceolate, and five nerved with the lateral nerves obscure. They are roughened and sometimes bear a short awn near the tip. Flowering occurs in June and July (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969).

Ecological Relationships and Distribution Figure 37—Prairie Junegrass panicles are spikelike Prairie Junegrass is the only species of Koeleria with short-appressed branches (photo courtesy of native to Western North America. It ranges from John Kinney, USDA FS, Rocky Mountain Research southern Ontario to British Columbia and southward Station, Boise, ID).

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1987, 1993; Cronquist and others 1977; Hitchcock or yellow starthistle (Borman and others 1991). In 1950; Hitchcock and others 1969). Plants are cold, heat, irrigated seed fields the plants grow relatively slowly and drought tolerant (USDA Natural Resources Con- compared to weeds (Smith and Smith 1997). Estab- servation Service 2003). lished plants are long lived (Hassell and others 1983). Prairie junegrass production is improved by spring or fall burns when adequate soil moisture is available, Uses and Management and effects may be long lasting (Blaisdell 1953). Late spring burns are most damaging (Towne and Owensby All classes of wildlife and livestock utilize prairie 1984). Plants are small and clumped with the growing junegrass with greatest use early in the season (Rose points near the soil surface; thus, they burn rapidly and others 1998). It provides good-quality early spring with little heat transfer (Young 1983). Recovery is forage due to its rapid development, but palatability from the crown or from seed. decreases during seed production (Vallentine 1961). Prairie junegrass is a key winter food source for Plant Culture animals of the British Columbian prairies (Blower 1982). Although utilized by whitetail and mule deer, Native stands of prairie junegrass are collected by prairie junegrass is not a primary source of forage hand stripping, while cultivated fields can be com- because of its scattered distribution. bined or swathed and combined (Smith and Smith Prairie junegrass can be used to revegetate highly 1997). Lodging and shattering are not generally prob- disturbed areas (Redmann and Schwarz 1986). Stud- lems with this species. The seeds are small, but readily ies of succession in the Piceance Basin of northwestern cleaned using an air-screen machine. Seed yields on Colorado (Biondini and others 1985) and Wood Buffalo irrigated land average 150 pounds per acre (168 kg per National Park, Alberta (Redman and Schwarz 1986), ha) and 75 pounds per acre (84 kg per ha) on showed that prairie junegrass increased following soil nonirrigated land (Smith and Smith 1997). There are disturbances. The species has been included in seed about 2,315,000 seeds per pound (5,090,000 per kg) mixes designed to restore disturbed fescue communi- (Smith and Smith 1997). Blake (1935) found viability ties in Glacier National Park (Laycock 1967). of seed kept in dry storage decreased from 21 percent after 3 years to 7 percent after 6 years (Blake 1935). Varieties and Ecotypes Prairie junegrass should be planted about 0.25 to 0.50 inch (0.6 to 1.3 cm) deep on a firm seedbed in fall. There are no releases. Seed fields and upper elevation sites are planted in early spring. Seed may be planted by drilling or broad- Oryzopsis hymenoides casting. Seed fields should be planted at 1 pound per acre (1.1 kg per ha) in 24-inch (60-cm) rows, 2 pounds Indian Ricegrass ______per acre (2.2 kg per ha) in 12-inch (30-cm) rows, and Synonyms 0.75 pound per acre (0.82 kg per ha) for 36-inch (90 cm) rows (Smith and Smith 1997). Direct seeding and Achnatherum hymenoides seedling transplanting has produced poor results in Stipa membranacea seed fields; however, transplanting 1-year-old plants Stipa hymenoides has been more successful (Nuzzo 1978). Commercial Oryzopsis membranacea growers do not grow junegrass often because they are unable to acquire large amounts of native seed (Hassell Description and others 1983). Available native seed is frequently of poor quality, as it is generally harvested from Indian ricegrass is a densely tufted perennial bunch- wildland sites. Use of locally adapted material may be grass with a deep, fibrous root system. Culms are essential for reestablishment of healthy stands hollow, thick walled, and 0.7 to 2.8 ft (2 to 8.5 dm) tall. (Simonin 2000). Some European material is now being Leaf sheaths are persistent and glabrous to minutely marketed in the United States. hairy. Old sheaths are fibrous and papery, sometimes Junegrass may be useful for reestablishing vegeta- becoming partly buried on sandy sites. Ligules are tion resistant to annual weed encroachment and inva- 0.10 to 0.30 inch (2.5 to 7.5 mm) long, entire to lacerate sion, largely due to the early spring emergence of its along the margins, and abruptly tapering at the apex. seedlings. This species does well in colonizing dis- Leaf blades are smooth, strongly involute, about turbed soils, and seedling emergence and survival are 0.04 inch (1 mm) wide, and nearly as long as the not affected by competition with Japanese brome (Romo culms. Panicles are 2 to 7.5 inches (5 to 19 cm) long and Eddleman 1987; Simonin 2000). However, plugs with the branches and hairlike branchlets paired, of prairie Junegrass were unable to compete with lacey, and spreading widely at maturity. Glumes are medusahead, dogtail, bulbous bluegrass, ripgut brome, ovate, fine hairy to nearly hairless, translucent along

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 393 Chapter 18 Grasses the margins, and tapering to a point or short awn up The growth cycle of Indian ricegrass is keyed to to 0.08 inch (2 mm) long. The three or sometimes five survival in dry and uncertain environments. Growth nerves are prominent near the base, but become indis- of mature plants is initiated at low temperatures in tinct toward the apex. Length of the first glume is 0.18 early spring (Pearson 1979). Leaves and flowers de- to 0.31 inch (4.5 to 8 mm), and the second is 0.17 to 0.30 velop rapidly, and good seed crops mature prior to inch (4.2 to 7.5 mm). Lemmas are 0.10 to 0.20 inch (2.5 summer drought. Flowering may continue into sum- to 5 mm) long, subglobose to fusiform, hard, thickened, mer in favorable precipitation years (Jones and Nielson shiny, and dark brown or nearly black when mature. 1989; Stoddart and Wilkinson 1938). Plants adapt to They are densely silky haired with the hairs nearly unstable sandy soils by developing adventitious roots exceeding the glumes. Awns are 0.12 to 0.22 inch (3 to from elongating basal internodes (Stoddart and 5.5 mm) long, straight, and readily deciduous. Paleas Wilkinson 1938). Fungal hyphae are associated with resemble the lemmas (Arnow 1987, 1993; Cronquist the root system (Robertson 1976), and the rhizosheaths and others 1977; Hitchcock 1950; Hitchcock and oth- fix nitrogen (Wullstein 1980). ers 1969). Plants flower from May through August. Indian ricegrass is fire tolerant when dormant. Spring Indian ricegrass is largely self-fertile, particularly in and early summer burns may be more damaging, but hot weather. Its chromosome number is 2n = 2x = 48 the effects of fire on the plant and its response to fire (Johnson and Rogler 1943). Indian ricegrass is the are not well documented. Recovery occurs through state grass of Utah and Nevada (USDA Natural Re- resprouting, tillering, or germination of seeds remain- sources Conservation Services 2002). It was named for ing in the soil seed bank or arriving from offsite its large, high protein seeds that were used by Native (Everett 1987; West 1994; Wright 1985; Wright and Americans to make flour. others 1979). Arnow (1987) transferred Indian ricegrass from Oryzopsis to Stipa based on similarities in embryologi- Plant Culture cal characteristics and floral structures and natural hybridization between the two genera. More recently, Rodent caching and burial of seeds by blowing sand Barkworth placed the species in Achnatherum provide appropriate microsites for natural regenera- (Barkworth and Everett 1987; Hickman 1993). tion (McAdoo and others 1983; Smigelski 1968). Long afterripening requirements may be met as seeds en- Ecological Relationships and Distribution dure drought periods, permitting development of stands during wet periods (Jones and Nielson 1989). Indian ricegrass is one of the most common grasses Seed production of native stands varies by year and on semiarid and arid lands of the West. It is distrib- is dependent upon local weather conditions. In addi- uted across Western North America from British Co- tion, flowering and seed ripening are indeterminate lumbia to southern California and Mexico and east- and seed dispersal is rapid (Robertson 1977). Conse- ward to Manitoba, the Dakotas, and Texas (Arnow quently, only a portion of the seed crop may be har- 1987; Stubbendieck and Jones 1996). It is one of the vested on any collection date. Narrow glume angle has most drought-tolerant native range grasses (USDA been used as a factor for selecting genotypes that Forest Service 1937), occurring on sand dunes and dry, might disperse seeds more slowly (Whalley and others sandy soils of valley bottoms where it often forms 1990). Much of the seed produced under arid condi- dense stands. Indian ricegrass is a dominant or asso- tions fails to fill, but remains on the plant; thus, ciated species in creosote bush, salt desert, big sage- production may be considerably less than anticipated brush, black sagebrush, pinyon-juniper, ponderosa (Stoddart and Wilkinson 1938; Stubbendieck and Jones pine, and mixed-grass prairie communities (Jensen 1996). In cultivated fields with irrigation, good seed and others 2001; Plummer and Frischknecht 1952; yields are expected annually. Young and Evans 1984). It also grows scattered among Seed may be harvested by hand collection from other vegetation on high grassy plains, foothills, ex- wildland stands or by swathing and combining (Wasser posed ridges, and dry, sandy or rocky mountain slopes. 1982). Planted seed fields are combined. Harvesting Indian ricegrass can be found at elevations ranging equipment such as brush harvesters for species with from 2,000 to 10,000 ft (600 to 3,000 m) (Booth and indeterminate flowering and fruit ripening may per- others 1980; Wasser 1982) on sites receiving 6 to 16 mit multiple harvests, thus increasing the total an- inches (15 to 40 cm) of annual precipitation (Natural nual yield (Stubbendieck and Jones 1996). Resources Conservation Service 2002). Although it Seed is cleaned using a barley debearder or generally occurs on sandy soils on drier sites, it is hammermill. Air-screen machines remove the chaff. found on soils ranging from sandy to heavy clays Seed is marketed at about 95 percent purity and 11 where precipitation is greater (Jones 1990; Platou and percent germination (Wasser 1982). A high percent- others 1986). age of the seed is usually dormant. There are about 159,000 seeds per pound (350,000 per kg) (Smith and

394 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

Smith 1997). Seed can be stored in a warehouse for 5 (Cook and others 1954; Stubbendieck and others 1985; years (Plummer and others 1968). Young 1989) (fig. 38). Protein content of dry material Emergence and establishment of seeded stands of is low during this period (Welch 1981), but some Indian ricegrass is slowed by complex seed dormancy material near the base of the plants may remain green (Jones 1990). Mechanical restraint imposed by the and more nutritious. The species is extremely impor- indurate lemma and palea restricts oxygen uptake. tant on winter ranges, such as salt desert shrublands Physiological dormancy is strongest in newly har- where extensive stands occur (Young 1989). It is also vested seed and decreases over time. Acid treatments valuable in late winter and early spring because it and mechanical scarification have been applied to initiates new growth sooner than many associated relieve mechanical dormancy, while use of old seed species (Green and others 1951; Hutchings and Stewart and moist prechilling treatments can reduce physi- 1953; Quinones 1981). Indian ricegrass is moderately ological dormancy. These treatments and combina- resistant to grazing, but stands have been lost as a tions of these treatments, however, have not proven result of excessive use (Rogler 1960; Stubbendieck and satisfactory for ensuring stand establishment (Jones others 1985). Deferment of livestock use in early 1990), and they are not easily applied on a large scale. spring may be required in some years to permit recov- Seed dormancy in this species is further complicated ery of plants that have been grazed excessively and to by the presence of seed polymorphism (Jones 1990). permit seed production and dispersal. Summer dor- The presence of two or more likely heritable seed types mant plants are more tolerant of use (Stubbendieck with differing germination requirements within a popu- and others 1985). lation serves as a natural survival mechanism. Seed Seeds of Indian ricegrass are large and high in polymorphism, however, also increases the difficulty protein. Big game, livestock, rodents, rabbits, birds, of establishing uniform seeded stands. and other small animals seek them out (McAdoo and Seeds should be fall planted in areas receiving pri- others 1989). Rodent gathering and caching of the marily winter precipitation to provide exposure to seeds aids in dispersal and establishment of the cool, moist winter conditions that serve to reduce species. physiological dormancy (Smith and Smith 1997). Seed Because it is an early successional species, Indian should be drilled 1 to 3 inches (2.5 to 7.6 cm) deep, or ricegrass is frequently used to revegetate disturbed broadcast and covered to this depth. Seed may be sites such as mine sites on sandy soils. It may be used planted up to 4 inches (10.2 cm) deep in very sandy alone or in mixtures. Although initial growth is slow, soils (Wasser 1982). If pure stands are drill seeded, 5 to 10 pounds of pure live seed per acre (5.6 to 11.2 kg PLS per ha) should be applied (Allison 1988; Pawnee Buttes Seed, Inc. 2002). The rate should be increased for broadcasting. Natural variation in dormancy among seeds and seed types within a population often trans- lates to erratic stand development. Early growth is slow, and good weed control is required in agricultural fields to permit establishment of a good stand. Wildland seedings should not be conducted in areas where dense stands of cheatgrass or other weeds are likely to create considerable competition. Indian ricegrass should not be drill seeded with highly competitive species. Stands should be protected from grazing until they are well established because younger plants are easily uprooted. Stands develop slowly, but they are persistent once established (Wasser 1982). Wildland seedings spread well from seed if not grazed excessively (Plummer and others 1968).

Uses and Management

Indian ricegrass is highly palatable to wildlife and Figure 38—Indian ricegrass is an extremely impor- livestock (Beale and Smith 1970; Kufeld 1973; Lauer tant forage species that remains nutritious through and Peek 1976). Plants are productive and the herbage the winter (photo courtesy of John Kinney, USDA cures well, providing a good energy source in winter FS, Rocky Mountain Research Station, Boise, ID).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 395 Chapter 18 Grasses established plants provide canopy cover and develop Phalaris arundinacea extensive root systems that bind surface soils in areas where few other species may be adapted. Indian Reed Canarygrass______ricegrass is used for low-maintenance landscaping in Synonyms recreational areas, on roadsides, and in parks and gardens. It is often used as an ornamental and in dried Arundo colorata bouquets because of its attractive growth habit and Calamagrostis variegata seed heads. Description Varieties and Ecotypes A perennial sodformer, reed canarygrass was first Seed production, seed germination, and establish- introduced into the United States in about 1880. By ment traits vary considerably among populations of the 1930s the first selections were released for use in Indian ricegrass. Specific ecotypes have been selected the Western States. The plant is now widespread and to provide cultivars that can be successfully cultured regarded as a widely naturalized species. Reed (Jones 1990; Jones and Nielsen 1989). The cultivars canarygrass grows to 9 ft (2.7 m) in height, and it is released to date are all nonbred, native-site popula- strongly rhizomatous. Leaf blades are flat and 0.24 to tions. 0.70 inch (6 to18 mm) wide. Ligules are 0.12 to 0.40 ‘Nezpar’ originated near White Bird, ID (Alderson inch (3 to 10 mm) long, blunt or rounded at the apex, and Sharp 1994), and has been seeded extensively in sometimes hairy, and usually lacerate along the mar- the West. It is best adapted to the Snake River Plain gins. Panicles are 2.8 to 7.9 inches (7 to 20 cm) long, and Columbia River Basin. compact, and sometimes lobed at the base. Spikelets ‘Paloma’ originated from a site near Pueblo, CO are ovate and three flowered with the first two florets (Alderson and Sharp 1994), at an elevation of 5,000 ft sterile and the third perfect. Glumes are subequal, (1,530 m) that receives 10 to 12 inches (250 to 300 mm) 0.16 to 3.0 inches (4 to 7.5 mm) long, and strongly of annual precipitation. Paloma is adapted to sandy to compressed, especially toward the tip. They are keeled sandy loam soils where it exhibits good stand establish- and three nerved, with the lateral nerves most promi- ment, vigor, drought tolerance, and forage production. nent. The keel of the glume is roughened, tapering to It is seeded to provide soil stabilization on mined lands a point, and usually wingless. Lemmas are shiny, and other disturbed rangelands. Paloma is adapted to appressed hairy, and faintly three to five nerved. The basin and Wyoming big sagebrush; black, Bigelow, and fertile lemma is 0.12 to 0.16 inch (3 to 4 mm) long, sand sage; and pinyon-juniper communities. It has ovate, and brownish at maturity. The two sterile performed well in Arizona, New Mexico, and the south- lemmas subtending the floret are subequal, 0.04 to ern portions of Utah, Nevada, and Colorado. 0.08 inch (1 to 2 mm) long, awl shaped, and hairy ‘Ribstone’ was collected near Taber, Alberta (Jones (Arnow 1987; Cronquist and others 1977). and Larsen, in press; Jones and others 2002e, 2004c), and selected for its good seed yield, germination, and Ecological Relationships and Distribution acute glume pair angle. The latter trait has been associated with seed retention. Ribstone is adapted to Reed canarygrass is a common riparian species in the dry mixed grass ecoregion characterized by brown wet places along ditches, streambanks, marshes, and chernozem soils in southeastern Alberta and sur- meadows (fig. 39). It ranges from Alaska through rounding areas. The Utah Agriculture Experiment Canada and the Western States to California, Nevada, Station recommended it for release as a Selected northern Arizona, northern New Mexico, and Okla- Germplasm in 2002. homa. It occurs in the Great Lakes area of the Central ‘Rimrock’ originated at 3,600 ft (1,100 m) near Billings, States and eastward to North Carolina. It is also found MT (Alderson and Sharp 1994; Jones and Larson, in in Mexico and South America. Reed canarygrass toler- press; Jones and others 2002e, 2004a), on a site with ates frequent and prolonged flooding and submer- fine sandy loam soil that receives 10 to 14 inches (250 gence, but it can also withstand periods of drought. It to 350 mm) of annual precipitation. It is adapted to is best adapted to fine-textured and poorly drained sandy soil, but it is more persistent in medium- and soils, and it is moderately tolerant of saline or alkaline heavy-textured soils than are Nezpar or Paloma. Rim- soils (Marquis and others 1984; Wasser 1982). It has rock is adapted to the Northern Great Plains, Rocky little shade tolerance. Mountains, Great Basin, Snake River Plain, Colum- Reed canarygrass has become a weedy species in bia River Plateau, and prairies of southern Alberta many wetland areas. It is highly productive and per- and Saskatchewan. Its intended uses are for revegeta- sistent once established. It spreads vegetatively and tion of disturbed rangelands and as a forage for live- from seed. It is highly competitive with timothy, Ken- stock, big game, birds, and other wildlife species. tucky bluegrass, and redtop, often invading areas

396 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

with degraded understories, and disturbed sites with poorly drained soils. It is a useful soil stabilizer and provides an abundance of forage for cattle and big game. Waterfowl, upland game birds, riparian mammals, and fish all use reed canarygrass for cover and food. Reed canarygrass hay fields have been planted to provide goose grazing areas (Burgess 1969). Prairie chickens use reed canarygrass for hiding and thermal cover in winter (Tester and Marshall 1962). Reed canarygrass has been rated as good forage for cattle, sheep, and horses in Utah, Colorado, Wyoming, and Montana (Dittberner and Olsen 1983). It is most palatable when it is growing, becoming more coarse and increasing in alkaloids in autumn (Boggs and others 1990). Reed canarygrass can be a desirable forage crop for cattle. Grazing should begin when the grass is 12 inches (30 cm) tall and when soils are dry to minimize trampling (Boggs and others 1990). Intense stocking rates with a short rotation period are recommended. Stands should not be grazed to less than 5 to 8 inches (13 to 20 cm) in height. Reed canarygrass is often cut Figure 39—Reed canarygrass, invasive in many for hay, providing an acceptable winter feed for cattle. North American wetland areas, forms monotypic This grass has been successfully established as an stands that displace native species (RMRS photo). understory with aspen and conifers. Although it has little shade tolerance, it can still dominate these communities. It has been seeded with other perennial grasses in aspen parks and mountain herblands, mead- dominated by these species and developing persistent, ows, and on wet sites, but it generally dominates the monotypic stands (Apfelbaum and Sams 1987). stand. Reed canarygrass has been widely seeded to control erosion and stabilize streambanks, as it pro- Plant Culture duces a spreading ground cover that provides excellent soil protection. Reed canarygrass has been seeded Reed canarygrass is highly productive and has been in the Intermountain West on areas with a high water successfully used to plant meadows, wetlands, and table, including sites that are inundated with water other sites with poorly drained soils. It can be seeded for up to 4 months. It has been planted within water- in fall or early spring when there is sufficient moisture ways and riparian areas from the valley floor up to and for establishment. Seeds of reed canarygrass germi- including the conifer forest. This grass will grow in nate readily, but produce weak seedlings. Conse- neutral to very basic salty soils. It has not proven quently, satisfactory stands often fail to develop. Prob- useful on most wildland plantings, and is now most lems arise due to the difficulty of preparing seedbeds widely planted as a pasture species at lower eleva- on wetland or wet meadow sites. Planting sites must tions. It has often been seeded in riparian communi- be allowed to drain or dry out prior to cultivation or ties because seeds of native species were not available, seeding. Soil crusting is common on these sites and but it gains dominance to the exclusion of other spe- contributes to poor seedbed conditions and planting cies. It should not be planted where native recovery is failures. Reed canarygrass should be seeded alone, as desired and wildlife habitat is important. it is not compatible with most other grasses. Once established, it reproduces mainly by rhizomes that Varieties and Ecotypes grow into dense mats within 1 year. Up to 74 percent of new shoots in established stands arise from rhi- ‘Ioreed’ reed canarygrass has been seeded in the zomes; new tillers bud within 2 inches (5 cm) of the soil Intermountain West on areas with high water tables, surface (Apfelbaum and Sams 1987). sites that are inundated for up to 4 months each year, and along waterways and riparian areas from the Uses and Management valley floor up through conifer forests. This grass will grow in neutral to very basic, salty soils. Ioreed was Reed canarygrass has been used to seed wetlands, selected for its high forage productivity and its adap- meadows, riparian sites, and aspen and conifer stands tation to wet sites. It is seeded to stabilize disturbed

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 397 Chapter 18 Grasses sites, waterways, and severely eroding areas, and to mud of bogs and marshes. It can also appear on provide forage. Ioreed can be very aggressive and relatively well-drained soils on grassy slopes, dry exclude other species. Other, less used, varieties in- meadows, and occasionally in moist sagebrush areas. clude ‘Castor’ and ‘Vantage’ (Alderson and Sharp In Alaska and along the northwest coast it grows at sea 1994). Seed of other unknown sources has been planted level. Alpine timothy may develop dense, monotypic widely in the Intermountain West. stands, but in many areas, it occurs mixed with other species, forming only a minor portion of the vegetation. Phleum alpinum Alpine timothy grows on deep clay loam to sandy Alpine Timothy, loam soils of pH 5.0 to 7.5 that are poorly to well Mountain Timothy ______drained (Hassell and others 1983), but it sometimes occurs on shallow soils in areas receiving 16 to 60 Synonyms inches (410 to 1,520 mm) of annual precipitation. Plants grow slowly and are moderately long lived Phleum pratense var. alpinum (USDA Natural Resources Conservation Service 2003). They reproduce from shoots developing from the base Description of the plants as well as from seed. The species is cold Alpine timothy is a densely tufted perennial with culms 0.7 to 1.5 ft (2 to 4.5 dm) tall that are often prostrate at the base. Leaf sheaths are glabrous. Leaf blades are flat, about 0.10 to 0.28 inch (2.5 to 7 mm) wide, roughened along the margins and sometimes on the surfaces. Auricles, when present, are blunt to rounded. Ligules are 0.02 to 0.18 inch (0.5 to 4.5 mm) long, truncate, and subentire. Panicles are small and ovoid to cylindric when fresh. When pressed flat they are 0.4 to 2 inches (1 to 5 cm) long and 0.28 to 0.47 inch (7 to 12 mm) wide (fig. 40). Spikelets are one flowered and elliptic. Glumes are subequal, 0.10 to 0.14 inch (2.5 to 3.5 mm) long, abruptly truncate, and tapering to stout awns that are 0.05 to 0.13 inch (1.2 to 3.2 mm) long. Margins of the glumes are roughened; the margin of the first glume is sometimes ciliate. Lemmas are 0.07 to 0.10 inch (1.7 to 2.5 mm) long, lanceolate to ovate, truncate at the apex, glabrous, and shiny or sometimes fine hairy. Alpine timothy is distinguished from timothy by the sheath of the uppermost leaf being typically inflated. Its panicles rarely exceed 2 inches (5 cm) in length, the awns are generally greater than 0.06 inch (1.5 mm) long, and the culms are not bulblike at the base (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969).

Ecological Relationships and Distribution Alpine timothy is the only species of Phleum native to North America. It is widely distributed throughout the cooler portions of North America and ranges from Alaska south to New Hampshire, northern Michigan, South Dakota, New Mexico, and southern California. It is also found in Mexico, Chile, Patagonia, Northern Europe, and Asia. Alpine timothy grows mainly in alpine and subalpine areas, and it also extends down Figure 40—Alpine timothy is the only Phleum species through the spruce-fir zone and into the aspen zone. It native to North America. Panicles generally less than 2 can be found in moist or wet mountain meadows, inches (5 cm) in length aid in distinguishing it from parks, on streambanks, around springs, and in the timothy (Hitchcock 1950).

398 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses and fire tolerant, fairly shade tolerant, but not drought 15 dm) tall. Leaf blades are flat and 0.12 to 0.31 inch tolerant (USDA Natural Resources Conservation Ser- (3 to 8 mm) wide with roughened margins and some- vice 2003). times with small auricles. Ligules are 0.06 to 0.20 inch (1.5 to 5 mm) long, subentire, rounded at the apex and Plant Culture sometimes lacerate along the margins. Panicles are 0.12 to 0.63 inch (3 to 16 mm) long, 0.20 to 0.40 inch (5 Supplies of alpine timothy seed are limited, as seed to 10 mm) wide when pressed flat, and compressed- is generally harvested from wildland stands. Seed cylindrical when fresh. Spikelets are one flowered and matures in August or September (USDA Forest Ser- elliptical. Glumes are subequal, 0.08 to 0.13 inch (2 to vice 1937) and is hand harvested (Rose and others 3.2 mm) long, abruptly truncate, and awned. Keels of 1998). Seeds remain on the plants for a short period the glumes are strongly and evenly ciliate. Glumes are before dispersal (USDA Natural Resources Conserva- three nerved, with the lateral nerves close to the keel. tion Service 2003). Fruits are cleaned using a The first glume is slightly narrower and sometimes hammermill and air-screen machine. Seeds are small hairy along the margins. Awns of the glumes are 0.02 at 1,680,000 seeds per pound (3,703,730 per kg) (Hassell to 0.06 inch (0.6 to 1.6 mm) long. Lemmas are 0.05 to and others 1983) and should be planted shallowly on 0.08 inch (1.2 to 2 mm) long, ovate, faintly five nerved, a firm seedbed in fall or spring by cultipacking or by minutely hairy, truncate, lacerate, and sometimes broadcasting. Seeds can be planted on or near the with a small awn at the apex (Arnow 1987, 1993; surface if the soil is moist. Germination and emer- Cronquist and others 1977; Hitchcock 1950; Hitchcock gence are good if moisture is available, but seedling and others 1969). growth is slow to moderate (Hassell and others 1983; Rose and others 1998). Ecological Relationships and Distribution Uses and Management Timothy is widely distributed in Europe and Asia in cool, humid habitats. Introduced to North America, it Alpine timothy produces a fair amount of nutritious is now widespread in the northern half of the United foliage that usually remains green throughout the States and southward in mountainous areas. It is summer, making it especially valuable as a late- found on moist meadows, weedy or grassy parks, season feed for livestock and big game (Hitchcock streambanks, moist canyon bottoms, open grassy 1950; Stubbendieck and others 1992). Regrowth fol- slopes, woodlands, forest openings, and roadsides and lowing use is slow (Natural Resource Conservation trails from sea level to 10,500 ft (3,200 m). Timothy is Service 2003). widely adapted, but grows best on deep, fertile, well- Alpine timothy spreads slowly from seed (Natural drained, moist clay, silt, or loam soils (Wasser 1982) Resources Conservation Service 2003). The species that are weakly basic to weakly acidic. It also grows in naturally revegetates disturbances and openings in sandy, gravelly, or rocky areas if the soil is moist. It mountainous areas (Brown and Johnston 1978a,b). It withstands winter and early spring flooding, but it is has performed well when tested on alpine mine wastes intolerant of flooding during the growing season in Montana. Alpine timothy has potential for use in (Jensen and others 2001). Timothy is not drought improving wildlife habitat, forage for livestock, road- tolerant, but it is cold and frost tolerant (USDA Natu- way plantings, and in seeding mixes for disturbed ral Resources Conservation Service 2003). Timothy is areas within its range. moderately fire tolerant, particularly when dormant. It occurs on many wet sites and high-elevation loca- Varieties and Ecotypes tions that rarely burn (Wasser 1982). There are no releases. Plant Culture Phleum pratense Timothy blooms in late spring and produces large numbers of seeds that mature in mid to late summer. Timothy ______Seeds are small and dispersed by livestock, wildlife, Synonyms wind, and other forces (USDA Forest Service 1937). Seed are harvested by combining, and are cleaned Phleum nodosum var. pratense with an air-screen machine. There are about 1,300,000 seeds per pound (2,865,980 per kg) (Wheeler and Hill Description 1957). Seeds remain viable for 4 to 5 years in dry storage (Wheeler and Hill 1957). They become highly Timothy is a tufted perennial with a usually bulbous germinable in about 1 month following harvest, and base, hairless leaf sheaths, and culms 0.7 to 4.9 ft (2 to

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 399 Chapter 18 Grasses germination remains high for 1 to 2 years. A potas- 1977; Hitchcock 1950; Hitchcock and others 1969). sium nitrate pretreatment and a 5-day prechill with Stands are short lived if not permitted to reseed incubation at 59/77 ∞F (15/25 ∞C) or 68/86 ∞F (20/30 ∞C) naturally. Established seedings are long lived when (16 hrs/8 hrs) and exposure to light during the high managed properly (USDA Natural Resources Conser- temperature periods are recommended for maximiz- vation Service 2003). ing germination (AOSA 1999). Germination occurs Timothy can be seeded where soils are moist and the rapidly. Seed lots may be purchased with 99 percent growing season is adequate for seed production. It has purity and 90 percent germination (Wasser 1982). been used for reseeding irrigated mountain meadows Timothy is seeded in fall where early spring drying and to provide ground cover and forage on burned, of the soil is likely to occur. Higher elevation sites are logged, or excessively grazed mountain rangelands planted in mid to late summer, while irrigated fields where annual precipitation exceeds 40 inches (102 cm) are planted in late summer (Wasser 1982). Seed should per year (Jensen and others 2001; USDA Forest Ser- be planted about 0.25 to 0.50 inch (0.6 to 1.1 cm) deep, vice 1937). An early to midseral species, timothy will or slightly deeper in coarser or drier soils (Jensen and encroach and dominate on disturbed sites. It is used in others 2001; Wasser 1982). Seed can be surface planted seeding mixes to stabilize roadsides, mined lands, or barely covered on fine, moist soils. The small seeds recreational areas, and other disturbed sites (fig. 41). may require addition of a diluent to permit regulation Timothy establishes by the second year after seeding. of the seeding rate. Seeding is accomplished by broad- Adapted seed sources should be used (Wasser 1982). casting, cultipacking, or use of a drill that will place Timothy can be invasive and limit the regeneration the seeds near the soil surface. and establishment of native species. It has reduced Timothy seedlings develop rapidly when adequate native grass colonization in fescue grasslands of Gla- moisture is available (Cornelius and Talbot 1955; cier National Park and invaded natural vegetation Hassell and others 1983; Jensen and others 2001; there (Tyser 1992). It competes strongly with seeded USDA Natural Resources Conservation Service 2003). and transplanted conifers where seeded following wild- They are susceptible to competition and soil drying fires (Baron 1962). because of their shallow root systems. Plants are generally fully established by the second growing Varieties and Ecotypes season, or sometimes by the end of the first growing season (Wasser 1982). Transplants with well-devel- ‘Climax’ is the most commonly seeded timothy on oped root systems are sometimes more successfully rangelands and wildlands in the Intermountain West. used in areas where frost heaving or drying are prob- It is adapted to aspen, ponderosa pine, conifer, and lems (Brown and Johnston 1978a,b; Brown and others subalpine communities. Climax is seeded to provide 1976). forage and soil stabilization. It can be seeded in fall or early spring. Germination occurs quickly, and seed- Uses and Management ling establishment and vigor are good. Climax is Timothy is known best for its widespread use as hay, primarily for horses, but it is also a forage plant where seeded on Western ranges. It is frequently grown in combination with other cool-season grasses and with red clover, alfalfa, and other legumes (Berg and others 1996; Wasser 1982). Timothy hay can be harvested two to four times per season, but regrowth is slow. The seed is used by birds. It provides cover for small mammals, waterfowl, and other birds. It is used as a forage by big game animals. Livestock use timothy in spring and fall, and it persists well if properly grazed (Wasser 1982). Timo- thy should not be grazed prior to emergence of the heads. Forage quality, quantity, and palatability are greatest in spring. Plants recover slowly following grazing, and forage quality declines rapidly as plants mature. Plants can spread by tillering, but they are Figure 41—Seeded on mountainous sites to pro- not resistant to heavy grazing or trampling because vide forage or soil stabilization, timothy limits estab- the sod and bulbous bases of the plants are easily lishment of naturally recovering or seeded species damaged (Arnow 1987, 1993; Cronquist and others (RMRS photo).

400 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses compatible with natives, providing rapid soil stabili- as Canada bluegrass and Kentucky bluegrass. As a zation as slower developing species establish and result, some specimens may have short rhizomes. mature. Stands weaken with time; their longevity is (Arnow 1987, 1993; Cronquist and others 1977; normally 5 to 9 years. Climax will reseed itself in weak Hitchcock 1950; Hitchcock and others 1969). Arnow communities and on open, disturbed sites if seed is (1987) placed Poa ampla in the P. secunda complex. allowed to mature. The variety, ‘Clair’, has been seeded recently. It Ecological Relationships and Distribution appears to perform as well as Climax on Western rangelands. Additional varieties developed for hay Big bluegrass is found on moderately moist to dry and pasture production for which seed is available are: sagebrush slopes, aspen and forest openings, and ‘Alexander’, ‘Alma’, ‘Argus’, ‘Basho’, ‘Bottnia II’, midelevations and, less frequently, subalpine mead- ‘Bounty’, ‘Carola’, ‘Champ’, ‘Drummond’, ‘Dynasty’, ows (Wasser 1982). It ranges from Alaska to North ‘Farol’, ‘Glenmor’, ‘Hokuo’, ‘Itasca’, ‘Korpa’, ‘Mariposa’, Dakota and south from northeastern California to ‘Mohawk’, ‘Nike’, ‘Richmond’, ‘Salvo’, ‘Tiiti’, ‘Tiller’, New Mexico (Arnow 1987, 1993; Cronquist and others ‘Toro’, and ‘Winmor’ (Alderson and Sharp 1994). 1977; Hitchcock 1950; Hitchcock and others 1969). Big bluegrass begins growth in early spring. At low elevations it matures in late spring or early summer, Poa ampla while at higher elevations or on sites with greater Big Bluegrass ______moisture availability, growth may continue later into the summer or throughout the season (Cronquist and Synonyms others 1977; Shaw and Cooper 1973; Wasser 1982). Big bluegrass grows in areas receiving 15 to 21 inches Poa secunda (381 to 533 mm) of annual precipitation, but it also Poa confusa occurs in areas receiving as little as 10 inches (254 mm) (Hassell and others 1983). Description Big bluegrass is adapted to a wide range of soil Big bluegrass is a robust, long-lived, densely tufted, textures, but it is generally found on clay loam to hairless to minutely hairy, green or blue-waxy peren- sandy loam soils that may be shallow and of low nial. Mature plants develop strong, fibrous root sys- fertility. It grows on soils with pH ranging from 6 to 8 tems. Culms are 1.3 to 5.9 ft (4 to 18 dm) tall with a (Hassell and others 1983), and it is moderately sensi- basal cluster of leaves 8 to 16 inches (20 to 40 cm) long tive to salinity. Plants are only weakly tolerant of early that generally wither early. Leaf sheaths are smooth spring flooding, high water tables, and poor drainage to roughened, generally shorter than the internodes, (Hassell and others 1983). Local ecotypes are tolerant and open less than one-fourth their length. Auricles of of cold or heat. The species is only moderately shade culm leaves are often hard and conspicuous. Leaf tolerant and not particularly drought tolerant. Plants blades are roughened along the margins, faintly blue- are fairly fire tolerant when dormant (Hassell and waxy, 0.06 to 0.14 inch (1.5 to 3.5 mm) wide, and others 1983) usually flat, but become involute as they dry. Ligules are short, thick, 0.04 to 0.08 inch (1 to 2 mm) long, and Plant Culture truncate or more often rounded. Panicles are narrow Seed fields are harvested using a combine or strip- and elongate, 0.31 to 0.87 inch (8 to 22 mm) long, per or by first windrowing (Archer and Bunch 1953). densely flowered, and usually pale green, but some- Seeds are cleaned in a fanning mill. Acceptable purity times tinged with purple. Spikelets are 0.22 to 0.43 is 90 percent and germination 70 percent (Wasser inch (5.5 to 11 mm) long, three to five flowered, and 1982). There are about 917,000 seeds per pound relatively narrow to somewhat cylindrical. The glumes (2,022,000 per kg) (Heady 1975). Seed requires a taper to a point, the first glume is 0.12 to 0.18 inch (3 prechill treatment of 7 days at 41 ∞F (5 ∞C) for germi- to 4.5 mm) long and the second 0.14 to 0.22 inch (3.5 to nation (Currie 1967; Haferkamp and McSwain 1951). 5.5 mm) long and about as long as the first floret. Seed can be stored for 5 years (Plummer and others Lemmas are 0.17 to 0.24 inch (4.2 to 6 mm) long, 1943). Seed should be planted 0.25 to 0.50 inch (0.6 to rounded at the apex, and usually rounded on the back 1.3 cm) deep on medium-textured soils. It can also be toward the base, becoming somewhat keeled towards broadcast and covered. Seedings should be conducted the tip. Lemmas are yellowish green with translucent in fall in drier areas. Spring or early summer seedings margins or sometimes brownish. They are usually may be successful on sites receiving greater annual hairless, but sometimes slightly hairy below. Flower- precipitation or in irrigated seed fields (Archer and ing occurs from May through July. Big bluegrass often Bunch 1953; Currie 1967; Wheeler and Hill 1957). hybridizes with rhizome-producing Poa species such

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 401 Chapter 18 Grasses

Seedling vigor is low to moderate and stands mature slowly. Seedlings are easily pulled up, particularly on wet or loose soils; thus, grazing should be avoided for two growing seasons or until the roots are securely anchored.

Uses and Management Big bluegrass is seeded on disturbed mountain big sagebrush sites and high mountain meadows (Jensen and others 2001). It is used for postfire seedings in forested areas. It is also seeded in hay fields and pastures with alfalfa and other grasses in both irrigated and nonirrigated fields. Such seedings may require weed control during the first season. Big blue- Figure 42—Sherman big bluegrass can be seeded grass may be seeded in mixtures with other native with other natives to provide wildlife forage and grasses, forbs, and shrubs to provide diverse native increase species diversity (photo courtesy of Scott vegetation and wildlife habitat. Established stands of Lambert, USDI BLM, Boise, ID). big bluegrass are persistent and competitive with cheatgrass and other weeds. Big bluegrass greens up early in spring. It can be grazed in spring or fall, and it is highly palatable during these periods. Palatability declines in early Poa compressa summer when moisture is inadequate. Fall rather than spring grazing and dense seedings that favor Canada Bluegrass ______vegetative development are approaches for minimiz- Synonyms ing pull-up problems resulting from grazing (Hyder and Sneva 1963). Grazing in spring should be delayed Paneion compressum until plants reach a height of at least 8 inches (20 cm). Plants are tolerant of grazing, but excessive use as Description plants are heading out can reduce growth during the subsequent year (Wasser 1982). Excessive grazing Canada bluegrass is an introduced, perennial, cool- and trampling can lead to loss of big bluegrass. season, rhizomatous sodformer that has naturalized Big bluegrass provides forage for wildlife and is used throughout much of the United States and Canada. by ungulates in spring and fall. Elk use it throughout Plants are wiry, erect, and bluish green throughout. the year. At higher elevations it provides summer Culms are 8 to 31 inches (2 to 8 dm) tall, strongly forage, remaining green throughout the season if ad- flattened, prostrate at the base, and often bent at the equate moisture is available and producing good fall nodes. Stems and leaf sheaths are characteristically regrowth. Small animals use big bluegrass as escape flattened. Leaf sheaths are open near the base, and so and thermal cover. Birds use it for nesting cover and strongly compressed as to appear winged. Leaf blades consume the seeds and foliage. are flat to folded, and 0.06 to 0.14 inch (1.5 to 3.5 mm) broad with short, boat-shaped tips. Ligules are to 0.02 Varieties and Ecotypes to 0.08 inch (0.5 to 2 mm) long, fairly stiff, more or less entire, and rounded to nearly squared at the apex. ‘Sherman’, collected near Moro, Sherman County, Panicles are 1.2 to 3.5 inches (3 to 9 cm) long, compact, OR (Alderson and Sharp 1994), is a large, long-lived dense, and usually narrow, but occasionally some- bluegrass that greens up very early as the snow melts. what open (fig. 43). Spikelets are 0.12 to 0.30 inch (3 to In the Intermountain West this variety is adapted to 7.5 mm) long, crowded, and usually three to seven mountain brush communities and openings in aspen flowered. Glumes are subequal and 0.07 to 0.13 inch and conifer forests. Sherman is used for soil stabiliza- (1.8 to 3.2 mm) long, the first one to three nerved, the tion, forage production, revegetation of disturbed lands, second broader and three nerved. Lemmas are 0.08 to and postfire seedings (fig. 42). Big game and livestock 0.12 inch (2 to 3 mm) long, firm, sometimes leathery, seek it out. It is nonaggressive and compatible with strongly keeled, more or less rounded, and often pur- other native species. plish at the tip. There are fine hairs on the keel and along the marginal nerves, especially toward the base.

402 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

tends to dominate only on soils that are too acid, droughty, or nutrient deficient for Kentucky bluegrass (Johnson and Nichols 1970).

Plant Culture Canada bluegrass reproduces from seed and rhizomes. Seeding is best accomplished in fall. There are more than 2.5 million seeds per pound (5.5 million per kg). Seed may be drilled or broadcast, but it should not be planted more than 0.12 inch (0.3 mm) deep, as light is required for germination (Fulbright and others 1982). Seedlings may emerge, but they do not persist with more competitive species on highly fertile soils. Seedlings do, however, compete exceptionally well on soil with low fertility (Sampson and others 1951), but they develop slowly, requiring 2 to 4 years to reach maturity. Once established, Canada bluegrass can increase its area of occurrence by natural spread, even amid considerable competition. In the Intermountain West, Canada bluegrass has been seeded in big sagebrush, mountain brush, aspen, conifer forests, mountain meadow parks, and riparian sites. It requires at least 14 inches (356 mm) of annual precipitation to Figure 43—Canada bluegrass panicles are establish and persist. Because seedlings develop slowly, compact, dense, and usually narrow, but sometimes open (photo courtesy of Kevin Jensen, Canada bluegrass can be seeded with tree and shrub USDA ARS, Logan, UT). species on roadways and other disturbances. Uses and Management Canada bluegrass has been used extensively as a The web is lacking, or if present, it is very scant (Arnow turf grass, especially in mixtures with other turf 1987; Cronquist and others 1977; Hitchcock 1950; species. It is often used to provide cover and erosion Hitchcock and others 1969). control on roadsides, road cuts and fills, borrow pits, dam sites, and recreational areas. It is frequently Ecological Relationships and Distribution seeded in mixtures with legumes for revegetation of mined areas (Hardy BBT Limited 1989). It is generally Canada bluegrass was introduced into North America slow to establish, but mature stands provide good from England. It is now widely distributed throughout cover and remain green over a long period. the cooler regions of North America from British Canada bluegrass is used by cattle, horses, sheep, Columbia southward to California and east to Geor- deer, and elk (Crawford and others 1969; Hardy BBT gia. In the United States, it is most common from New Limited 1989; Kufeld 1973; Kufeld and others 1973; England to Ohio and West Virginia. It occurs sparsely Sampson and others 1951). It greens up early in throughout Intermountain ranges in mountain mead- spring, and growth continues in summer and fall. ows and along dry streambanks, and it is occasionally Consequently, it is palatable to livestock during this abundant on sites where it has been seeded. In the period (Hardy BBT Limited 1989). Because it matures Southwest it occurs on higher mountain ranges. It has later than Kentucky bluegrass and is more drought spread to occupy extensive disturbances throughout tolerant, Canada bluegrass tends to be more palatable its present range (Van Dyne 1958), and also grows in in summer. It provides fair to good cover for birds and irrigated lawns and pastures. small mammals (Dittberner and Olson 1983). This Canada bluegrass grows on soils ranging from clay species is resistant to heavy use and trampling and to sand and gravel. It grows well on soils with low shows good fire tolerance. fertility or poor drainage (Hardy 1989; Sampson and Canada bluegrass is classed as an invader on over- others 1951). Plants exhibit moderate drought and grazed rangelands (Van Dyne and Payne 1964). It is salinity tolerance, but they are not shade tolerant generally not recommended for seeding as a pasture (Van Dyne 1958). Canada bluegrass and Kentucky grass because of its low productivity. On poor soils, bluegrass occur on similar sites. Canada bluegrass mine wastes, and roadway disturbances it may be

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 403 Chapter 18 Grasses seeded to provide useful forage (Stubbendieck and involute with keeled tips. Culm blades are reduced or others 1985; Welsh and others 1987) and ground lacking, revealing only the ligule on the bladeless cover. Although slow to establish, this species com- sheath. Ligules are highly variable, often truncate, petes well with annual weeds. It is not recommended and range from 0.02 to 0.47 inch (0.5 to 12 mm) in in seedings where native species are desired. It can length to much elongate and tapering, sometimes be planted to furnish cover on harsh sites, but it exceeding 0.4 inch (1 cm) in length. Most are about usually does not support establishment of herba- 0.12 to 0.31 inch (3 to 8 mm) long. Panicles are pale to ceous species. Native trees and shrubs are able to deep purple or sometimes tawny, 0.12 to 0.6 inch (3 to establish into existing stands of the grass, particu- 15 mm) long, and 0.4 to 0.8 inch (1 to 2 cm) wide, larly in areas receiving over 20 inches (508 mm) of oblong, contracted, and usually situated well above annual precipitation. the basal leaves. Spikelets are 0.20 to 0.41 inch (5 to 10.5 mm) long and tightly three to six flowered. They Varieties and Ecotypes are strongly compressed, keeled throughout, and often appear papery. The first glume is 0.11 to 0.18 inch (2.8 ‘Foothills’ Selected Germplasm, a composite of eight to 4.5 mm) long; the second is 0.14 to 0.22 inch (3.5 to Canada bluegrass accessions (Englert and others 2002), 5.5 mm) long. Lemmas are 0.16 to 0.28 inch (4 to 7 mm) is adapted to the Northern Rocky Mountains of the long, blunt on the usually irregularly lacerated apex, United States and Southern Canada. It was selected and with long, silky hairs on the prominently nerved for its forage production, seed production, and ability keel and marginal nerves. Specimens of mutton blue- to spread. Recommended uses include erosion control, grass from the Intermountain area are generally pis- critical area plantings, grazing, and low maintenance tillate, posing doubt that fertilization actually takes landscaping. place. Some plants produce a few staminate or perfect ‘Reubens’ originated near Reubens, ID (Alderson florets (Arnow 1987) (fig. 44). Plants flower from May and Sharp 1994). It was selected for stabilization of low-fertility soils with an irregular moisture supply. In the Intermountain West it is seeded extensively in mountain brush, mountain big sagebrush, three-tip sagebrush, and ponderosa pine communities. Reubens establishes well on sterile, coarse-textured soils ranging from slightly acidic to slightly basic. It is seeded on mined lands, road cuts and fills, and other disturbances to stabilize the soil. Plants are low growing and not highly productive. Reubens is compatible with other species, and can be seeded as a component of mixtures.

Poa fendleriana Mutton Bluegrass ______Synonyms Eragrostis fendleriana Poa eatoni Poa montana Poa longiligula

Description Mutton bluegrass is a perennial bunchgrass that forms large tufts 0.7 to 2.5 ft (2 to 7.5 dm) tall. Stems tend to stool at the edge of the tufts, and there are occasionally a few short rhizomes. Leaves are basal, erect, numerous, and strongly blue waxy. Basal leaf sheaths are papery, persistent, somewhat roughened, Figure 44—Mutton bluegrass flowers from May and open nearly their entire length. Leaf blades are to August. Flowers are generally pistillate, but firm, stiff, usually roughened, 0.06 to 0.16 inch (1.5 to some plants produce a few staminate florets 4 mm) wide, sometimes flat, but more often folded or (RMRS photo).

404 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses to August (Arnow 1987, 1993; Cronquist and others Plant Culture 1977; Hitchcock 1950; Hitchcock and others 1969). Muttongrass forms a highly polymorphic complex Muttongrass seed matures between late May and extending across a broad elevational range and occu- mid-August (Sampson 1924; Smith and Smith 1997). pying diverse habitats (Arnow 1987). Soreng (1985) Seeds can shatter rapidly at maturity (Smith and described the species as including three subspecies Smith 1997). Seed stocks remain erect at maturity and differing in production of rhizomes, ligule characteris- extend well above the leaves. Fields can be direct tics, and reproductive biology. Goodrich and Neese combined as lodging is generally not a problem. Yields (1986) consider the species highly variable, but sug- in irrigated seed fields vary considerably from year to gested the subspecies be reduced to varieties. Al- year, averaging about 35 pounds per acre (39 kg per though most plants are pistillate, they found a limited ha) and ranging from 25 to 75 pounds per acre (28 to number of staminate plants that produced long 84 kg per ha) (Smith and Smith 1997). rhizomes. Seeds can be cleaned to some extent during combining. Further processing using a debearder may be Ecological Relationships and Distribution required to complete separation of seed from the debris. Trash is removed with an air-screen machine. Mutton bluegrass is one of the most widely distrib- There are about 890,000 seeds per pound (1,958,000 uted native bluegrass species in the West (Wedin and per kg). Germination tests can be conducted by incu- Huff 1996). It is distributed from southeastern British bating seeds at an alternating temperature regime of Columbia to Manitoba and south to California, North- 68/86 ∞F (20/30 ∞C) (16 hrs/8 hrs) with exposure to light ern Mexico, western Texas, Colorado, and western during the high-temperature alternation (Haferkamp South Dakota. In the southern part of its range it and McSwain 1951). Seed can be kept in dry, cold occurs from the pinyon-juniper zone through oakbrush, storage for up to 5 years (Smith and Smith 1997). mountain brush, ponderosa pine, and aspen commu- Mutton bluegrass seed fields can be planted in early nities to elevations of 7,000 to 12,000 ft (2,134 to spring. Establishment may require 2 years, and weeds 3,658 m) on sites receiving 10 to 22 inches (254 to must be controlled to reduce competition. Stand life is 559 mm) of annual precipitation (Link 1993; USDA about 8 years (Smith and Smith 1997). Natural Resources Conservation Service 2003). It ap- Wildland seedings should be conducted in fall. Spring pears mainly on mesas, ridges, and slopes, and in open plantings may be attempted, as emergence can occur timbered areas, as well as in well-drained parks and rather rapidly. However, spring plantings can be lost meadows. In the northern portion of its range it occurs when soil surfaces dry quickly in spring. Seed can be at lower elevations in the sagebrush zone where it planted by drilling if seeds are placed near the soil grows with other cool-season grasses and forbs. It surface. Seed may also be broadcast and covered or occasionally occurs in desert shrub communities seeded with a cultipacker. Mutton bluegrass seedlings (Arnow 1987), but it may have been lost from drier are generally quite small when they enter dormancy areas as a result of overgrazing. during the first growing season, but they begin growth Mutton bluegrass is best adapted to clay loam soils, early in the second growing season. Growth rate and but will grow on well-drained, medium- to coarse- lifespan are both moderate (USDA Natural Resources textured soils that are fairly neutral (USDA Forest Conservation Service 2003). Established stands spread Service 1937). In oakbrush, mountain brush, and slowly from seed, and generally do not interfere with aspen communities it grows as an understory species the establishment of other native species. on deep soils and tolerates some shade. In sagebrush and pinyon-juniper communities it may occur on shal- Uses and Management low and gravelly soils in addition to soils with moderately well-developed surface horizons (USDA Forest Mutton bluegrass begins growth in early spring and Service 1937; Dittberner and Olsen 1983). is grazed by livestock and wildlife. Palatability de- Mutton bluegrass is an important late successional creases as the plants mature, but they retain some species in many shrub-dominated communities (Koniak green leaves and are grazed late in the season when 1985). It is generally tolerant of fires (Arnold 1950). other forage is not available. Plants normally green up Although Koniak (1985) found some stands were dam- in the fall and offer good-quality forage at this period. aged by wildfires, Gartner and others (1978) deter- Mutton bluegrass persists well in openings and in the mined that growth of rhizomatous population was understory with oak and other shrubs, furnishing actually stimulated by fire. Plants have responded green forage over a long period. It is also an important well following fire in sagebrush, pinyon-juniper, and species within the sagebrush zone. Mutton bluegrass mountain brush woodlands in central and eastern is considered an excellent species for all classes of Utah. domestic livestock. It provides forage for deer and elk

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 405 Chapter 18 Grasses in spring, and is also used in winter when the foliage inch (2 to 3 mm) wide, spikelets mostly three flowered, is dry (Judd 1962). Pronghorns use the seedheads in and lemmas heavily webbed at the base. winter (Ferrel and Leach 1950). Mule deer and elk use Kentucky bluegrass is a leafy, long-lived perennial stands of this species in areas where pinyon and that spreads by seed and especially by rhizomes. It can juniper trees have been cleared. These stands can form a dense sod, especially on moist, fertile soils, even draw game animals from nearby cultivated fields, when grazed or clipped. Most roots are within 3 inches thus reducing crop damage. Stands can, however, be (7.5 cm) of the soil surface. Leaf sheaths are smooth, reduced by excessive grazing. The seeds and foliage occasionally somewhat roughened, and closed for about are used by birds (Christensen 1958; Harper and half their length. Blades are soft, usually green or others 1958; Martin and others 1951). sometimes slightly bluish, flat or more commonly Mutton bluegrass produces a fibrous root system folded, and about 0.04 to 0.18 inch (1 to 4.5 mm) wide about 10 inches (25.4 cm) deep that provides good with roughened margins and boat-shaped tips. Ligules surface erosion control. Like Sandberg bluegrass, this are shorter than wide, squared, and mostly entire. The species is drought tolerant. Its growth rate and lifespan inflorescence is an open, pyramidal panicle, 2 to 8 are both moderate (USDA Natural Resources Conser- inches (5 to 20 cm) long. Panicle branches are spread- vation Service 2003). Seed crops vary among years, ing or ascending and whorled in groups of three to five. and natural spread can be slow. Good recovery has Spikelets are 0.12 to 0.24 inch (3 to 6 mm) long, ovate, been observed on sites protected from grazing. congested, and two to four flowered. They are green to Mutton bluegrass has not been widely used to re- purplish in color and strongly compressed. Glumes are store disturbed areas, but it could be useful in efforts roughened on the keel; the first glume is one nerved to reestablish native diversity. The species could be and the second is three nerved. Lemmas are three to used in many areas where introduced grasses are now five nerved, with weak lateral nerves. They are rounded planted. It is key in areas where pinyon and juniper or tapered to a point apically and there are cobweb-like encroachment has depleted the understory and sea- hairs at the base (Arnow 1987; Cronquist and others sonal habitat for big game has been lost. It is also an 1977; Hitchcock 1950; Hitchcock and others 1969; important species to develop for use in seeding big USDA Forest Service 1937). sagebrush communities where the native understory has been lost and annual weeds have become a major Ecological Relationships and Distribution problem. Kentucky bluegrass was one of the earliest grass Varieties and Ecotypes introductions to arrive in the United States. Plantings in North America were reported as early as 1685 There are no releases. (USDA Forest Service 1937). Kentucky bluegrass is now widely distributed throughout most of North America, growing in every State and Canadian Prov- Poa pratensis ince. It is adapted for growth in cool, humid climates, Kentucky Bluegrass ______and is most prevalent in the Northern United States and Southern Canada. It is not common in the Gulf Synonyms States nor in the desert regions of the Southwest Paneion pratense (Wheeler and Hill 1957). Kentucky bluegrass is not Poa peckii widely adapted to arid regions receiving less than16 Poa agassizensis inches (40 cm) of annual precipitation. The grass, however, has spread into many wildland communi- Description ties. Kentucky bluegrass can be found on prairies and in cultivated fields, mountain grasslands, mountain Kentucky bluegrass is considered a European intro- brush communities, mountain meadows, riparian duction by many botanists, but its occurrence in re- drainages, and open forests. It is common along road- mote areas throughout the Intermountain region sug- sides and in waste places. gests that there may be native populations. Some Kentucky bluegrass has proven well adapted to publications treat Poa agassizensis as the native coun- many mountain brush, aspen and conifer communi- terpart of Kentucky bluegrass. They are differentiated ties, and wet or semiwet meadows in aspen and subal- as follows: Poa agassizensis has waxy-blue leaves with pine communities. It dominates riparian areas over a blades 0.03 to 0.08 inch (0.8 to 2 mm) wide, spikelets wide range of elevations. Considerable variability has mostly two flowered, and lemmas only slightly webbed been noted among collections of this species. Although as opposed to highly webbed at the base. Kentucky Kentucky bluegrass is generally best suited to areas bluegrass has bright green leaves, blades 0.08 to 0.12 receiving more than 16 inches (40 cm) of annual

406 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses precipitation, it has sometimes persisted when seeded is often abundant in mountain grasslands, moist and in more arid sites. Vallentine (1961) noted that the dry mountain meadows, aspen parklands, open pon- species is not shade tolerant, but numerous plantings derosa pine forests, and in riparian areas where it in forested communities have survived. Although Ken- receives heavy use by domestic sheep and cattle (Bowns tucky bluegrass grows best in full sunlight, it will and Bagley 1986; Clary 1975a; Kauffman and others tolerate light shading if moisture and nutrient condi- 1983). Mountain meadows dominated by Kentucky tions are favorable (Hardy BBT Limited 1989; Smoliak bluegrass may be relatively limited in extent, but they and others 1981). are highly productive and provide substantial amounts Kentucky bluegrass does not produce vigorous seed- of summer forage (McInnis and Vavra 1986). Plants lings. Plants persist once established, but stands are are highly resistant to grazing because their growing often slow to develop. Once established, however, the points remain below ground throughout the growing species can be aggressive, spreading by development season, and they produce a low ratio of reproductive to of strong rhizomes. It often invades and suppresses vegetative stems (Ehrenreich and Aikman 1963). Few populations of other useful species. Kentucky bluegrasses withstand heavy grazing as well as Kentucky grass produces abundant seed that establishes in bluegrass. It increases rapidly on overgrazed pastures openings within the community. It often dominates in and ranges, and its presence is usually an indication of heavily grazed meadows, pastures, riparian sites, and poor grazing management. For livestock use, these other disturbances. sites are best managed under a grazing system other Kentucky bluegrass grows on a wide variety of soils, than season long. but thrives on well-drained loams or clay loams rich in Kentucky bluegrass is highly palatable to most large humus (USDA Forest Service 1937), including soils ungulates in spring when it is green and succulent. In derived from limestone (Hardy BBT Limited 1989; moist mountain meadows, palatability remains mod- Smoliak and others 1981; USDA Forest Service 1937). erately high during summer (Bellrose 1980). In drier Optimal soil pH is between 5.8 and 8.2 (Smoliak and areas it becomes semidormant in summer and its others 1981). This species requires fertile soils and palatability is much reduced. Regionally, Kentucky exhibits a high nitrogen requirement at all stages of bluegrass is important in the diets of elk, mule deer, growth. and bighorn sheep (Dittberner and Olson 1983). Grass- lands dominated by this grass provide habitat for Plant Culture numerous small mammals (George and others 1978; Medin and Clary 1989). Kentucky bluegrass meadows There are 2.1 to 2.2 million seeds per pound (4.6 to found along mountain streams are often preferred 4.8 million per kg). Germination is generally high, foraging areas for wild ungulates and grouse. Mueggler ranging from 75 to 94 percent. Seed has been drilled or and Campbell (1986), however, suggested that the broadcast seeded in fall, but it should not be planted aspen/Kentucky bluegrass community type in Utah is more than 0.20 inch (5 mm) deep because exposure to one of the poorest aspen types for wildlife habitat due light is required for germination (Fulbright and oth- to its low diversity of plant species. ers 1982). If used, this species should be seeded in Kentucky bluegrass leaves and seeds are eaten by limited amounts in mixtures with other species due small mammals and songbirds (Martin and others to its competitive ability. Kentucky bluegrass estab- 1951). Prairie chickens eat small amounts of its seeds lishes well from broadcast seedings on exposed soils. It (Crawford and others 1969). It is also an important has been widely seeded by aerial application on range- food for cottontail rabbits and wild turkeys (Crawford lands and over mountainous terrain. It is also broad- and others 1969; George and others 1978). Sage- cast seeded on loose soils, including mine spoils and grouse use Kentucky bluegrass-dominated areas for roadway disturbances. Mine wastes and other exposed strutting and chick rearing. substrata must be fertilized to assure seedling estab- The use of Kentucky bluegrass for revegetation on lishment. Seeded stands are slow to develop. Once rangelands is limited because of its slow establish- established, however, Kentucky bluegrass is highly ment, lack of drought tolerance, and its high soil persistent and spreads from seed and rhizomes. fertility requirements (Vogel 1981; Wasser 1982). When planted in seed mixtures, it often takes 2 to 3 years to Uses and Management become established (fig. 45). The value of Kentucky bluegrass on rangeland is Because of its sodforming ability, Kentucky blue- also limited because of its low productivity, summer grass has been widely used as a ground cover. It is one dormancy, and propensity to invade native grass- of the first grasses to resume growth in late winter and lands. In the Intermountain West, Kentucky blue- to green up in early spring. It is highly palatable in the grass is well adapted to meadows with seasonally high early stages of growth and provides nutritious forage water tables and midsummer drought (Volland 1985a). for all classes of livestock and wildlife. In the West, it It has become naturalized and dominates many

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 407 Chapter 18 Grasses

Newport is rhizomatous, moderately drought tolerant, and exhibits good growth compared to other Kentucky bluegrass accessions. Newport is less competitive than some Kentucky bluegrass varieties, and it can be seeded in mixtures with some species.

Poa secunda Sandberg Bluegrass ______Synonyms Paneion sandbergii Poa canbyi Poa confusa Poa nevadensis Poa sandbergii Figure 45—Two-year-old Kentucky bluegrass plants Poa scabrella spread rhizomatously and develop a dense ground cover (RMRS photo). Description The Poa secunda complex occupies a wide range of habitats, and its members exhibit many forms, several having been given species status over the years. meadows once occupied by tufted hairgrass and sedges. Sandberg bluegrass is one of the more common early- Reestablishment of the original natives is impractical season desert bunchgrasses in the Intermountain area. because of the competive ability of Kentucky blue- It is a small, densely tufted perennial that often grass. produces extensive tufts of basal leaves that wither Because of its shallow root system, Kentucky blue- and disappear early (fig. 46). Plants are often more or grass is not recommended as a soil stabilizer. It is less less purple tinged throughout. Culms are 0.7 to 1.5 ft effective than the native grasses and forbs it has (2 to 4.5 dm) tall and wiry. Sheaths are smooth to replaced, and it is not capable of stabilizing roughened and open for three-quarters or more of streambanks. Erosion and channel downcutting may occur when it dominates, especially in heavily grazed areas (Hansen and others 1988a,b; Kovalchik 1987). Kentucky bluegrass is intolerant of prolonged flooding, high water tables, and poor drainage (Wasser 1982). Kentucky bluegrass is one of America’s most popular lawn grasses. It withstands considerable abuse and it is often used as a sod grass on golf courses, campgrounds, and ski slopes (Shaw and Cooper 1973). It fails to control weed invasion and spread, particularly in riparian sites and meadows. Serious weed problems occur where this species has displaced native grasslike species.

Varieties and Ecotypes Many varieties of Kentucky bluegrass have been selected and released for turf purposes. Most are not adapted for rangeland use. The most popular variety for wildland seeding is ‘Newport’, obtained from a Figure 46—A small, tufted cool-season perennial, coastal site in Lincoln County, OR (Alderson and Sandberg bluegrass is one of the most common Sharp 1994). Newport is seeded in the Intermountain bunchgrasses of the Intermountain area (photo cour- West to provide soil stabilization. It is adapted to tesy of Loren St. John, USDA NRCS, Aberdeen Plant well-drained alluvial soils derived from limestone. Materials Center, Aberdeen, ID).

408 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses their length. Leaf blades are soft, folded or involute, rooting system provide stabilization of surface soils rarely flat, 0.02 to 0.06 inch (0.5 to 1.5 mm) wide, and (Jensen and others 2001). 1.2 to 1.95 inches (3 to 5 cm) long. The basal cluster of Sandberg bluegrass is generally not damaged by leaves is 1.2 to 5.5 inches (3 to 14 cm) tall. The leaves wildfires. The small clumps, low amounts of litter, and are roughened and their tips are prow shaped. Ligules prolonged summer dormancy preclude extensive dam- are tapered to a point and rather prominent. Those of age to the buds. Early fires during dry years may be the basal leaves are mostly less than 0.06 inch (1.5 more damaging (Kearney and others 1960). Cover mm) long, and those of the culm leaves are usually provided by the species may increase for a time follow- more than 0.10 inch (2.5 mm) long. Panicles are 0.8 to ing fire if other species are lost or reduced (Blackburn 7.9 inches (2 to 20 cm) long, narrow to rather open, but and others 1971). Reestablishment of sagebrush or usually not dense. Panicle branches are short and other taller species may eventually reduce the cover usually ascending. Spikelets are 0.16 to 0.37 inch (4 to provided by Sandberg bluegrass, as it is not shade 9.5 mm) long, two to six flowered, slightly compressed tolerant (Acker 1992; Tueller 1962). and rather cylindrical, narrow, and pointed. They are usually strongly purple tinged. Glumes are rough- Plant Culture ened, translucent along the margins, and tapered to a point. The first glume is 0.09 to 0.20 inch (2.2 to 5 mm) Regeneration of Sandberg bluegrass is by seed and long and one nerved; the second is 0.12 to 0.20 inch (3 tillering with occasional production of a small number to 5 mm) long and three nerved. Lemmas are 0.12 to of rhizomes (Arnow 1987). The species is self-fertile or 0.22 inch (3 to 5.5 mm) long, rounded below and only wind pollinated and facultatively apomictic slightly keeled above. They become increasingly curly (Daubenmire 1970; Hitchcock and others 1969; Kellogg haired below, and are not at all webbed at the base 1985a,b). Plants produce considerable seed in early (Arnow 1987, 1993; Cronquist and others 1977; summer. Native stands are harvested by hand or with Hitchcock 1950; Hitchcock and others 1969). mechanical strippers. Little seed is normally harvested from wildlands, and such seed is often of low Ecological Relationships and Distribution viability. Seed fields of the released varieties are harvested by swathing (Smith and Smith 1997) or by Sandberg bluegrass ranges from Alaska through direct combining (Rose and others 1998). Some seed much of Canada and south into the Western United may be lost through shattering. Seed is cleaned using States, Mexico, Nebraska, and Michigan. The species a hammermill and air-screen machine. is also found in South America. It grows at elevations There are about 925,000 seeds per pound (2,039,255 from 980 to 12,000 ft (300 to 3,660 m) on sites receiving per kg) (Hafenrichter and others 1968). Yields from 10 to 22 inches (254 to 559 mm) of annual precipitation seed fields of the Canby bluegrass type average 150 (Hitchcock 1950; Stubbendieck and others 1992; USDA pounds per acre (168 kg per ha) (Smith and Smith Forest Service 1937; USDA Natural Resources Con- 1997). Wasser (1982) recommended 90 percent purity servation Service 2003). Sandberg bluegrass occurs on and 65 percent germination as purchasing standards. dry to mesic plains, foothills, and ridgetops of salt AOSA (1999) recommendations for germination are desert shrub, sagebrush, pinyon-juniper, oakbrush, pretreatment with 0.2 percent potassium nitrate fol- mountain brush, grassland, aspen, conifer, and meadow lowed by incubation at alternating temperatures of 68 communities (Arnow 1987). and 86 ∞F (20 and 30 ∞C) (16 hrs/8 hrs) with exposure Sandberg bluegrass occurs on soils of a wide range of to light during the high-temperature periods. textures, but it is most productive on deep, well- Seed fields and wildland seedings of Sandberg blue- drained sandy clay, silt, and sandy loam soils that grass should be planted on firm seedbeds in clay loam range from slightly acidic to basic. It is often a domi- to sandy loam soils at a 0.25 to 0.5 inch (0.6 to 1.3 cm) nant species on dry, rocky sites and on shallow soils of depth in fall (Fulbright and others 1982). Germination scablands. The species is tolerant of salinity and of is slow and seedling vigor is low. Stands establish sites that are flooded in spring, but it does not tolerate slowly and there is little production during the first soils that are saturated for prolonged periods (Arnow season (Smith and Smith 1997; Wasser 1982). Stand 1987; Hafenrichter and others 1968; Plummer and life for seed fields is 7 or 8 years. Use of local seed others 1968). sources is recommended for wildland plantings be- Sandberg bluegrass begins growth in early spring cause of the wide distribution of the species. prior to most other native bunchgrasses and sets seed in late spring or early summer, depending upon eleva- Uses and Management tion. Plants evade summer drought by entering dormancy. Growth may resume in fall if adequate precipi- Sandberg bluegrass has been successfully seeded in tation occurs. The dense tufts of leaves and the fibrous areas receiving as little as 12 inches (300 mm) of

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 409 Chapter 18 Grasses annual precipitation (Arnow 1987). Seedlings are not Sitanion hystrix highly competitive. New seedings must be protected from grazing and weed invasion. Established plants Squirreltail, bottlebrush ______are more competitive, but also compatible with other Synonyms native species. Plants are short lived; management should permit periodic seed production and natural Elymus elymoides seeding to perpetuate stands and reduce the risk of Aegilops hystrix cheatgrass invasion. Seedling establishment and stand Sitanion elymoides productivity fluctuate widely with weather conditions Sitanion californicum (Daubenmire 1975b). Sitanion rigidum Sandberg bluegrass is one of the earliest grasses to Sitanion glabrum be grazed in spring. It provides high-quality and Sitanion cinereum palatable, but coarse, forage for livestock and wildlife Sitanion montanum from spring to early summer. Both forage quality and Sitanion brevifolium palatability decline rapidly as the plant matures, but Sitanion longifolium the species receives some use in summer and fall. Sitanion hordeoides Summer regrowth is limited; fall regrowth may be moderate, depending upon precipitation (Thornburg Description 1982). Plants are highly resistant to grazing and trampling. Squirreltail is a short-lived, loosely to densely cae- Sandberg bluegrass is an important component of spitose perennial that is nearly hairless to white hairy the Townsend ground squirrel’s diet (Johnson 1977). throughout. Culms are erect to spreading and 0.3 to It provides some cover for small mammals, songbirds 2.1 ft (1 to 6.5 dm) tall, with hairless to silky haired leaf and waterfowl, and other small animals. Elk, deer, sheaths. Leaf blades are flat to folded or involute, 0.4 and other wildlife species browse it (Vallentine 1967). to 0.24 inch (1 to 6 mm) wide, stiff haired to soft hairy Sandberg bluegrass provides some surface soil stabi- above, and sometimes hairless below. Auricles are lization, particularly on shallow soils where it may be inconspicuous, up to 0.04 inch (1 mm) long, and often the dominant species. purplish. Ligules are short, less than 0.2 inch (5 mm) long, and squared at the margins. Spikes are compact Varieties and Ecotypes to loose and open, and 1.2 to 5.9 inches (3 to 15 cm) long, excluding the awns. They are often included in ‘Canbar’ originated in the Blue Mountains of the upper leaf sheaths, or sometimes entirely exerted Columbia County, WA (Alderson and Sharp 1994). It in robust specimens. The rachis disarticulates as the is adapted to basin and Wyoming big sagebrush and spikelets mature. Spikelets are mostly in pairs, some- mountain brush communities. Although it estab- times in threes, or solitary when in the upper portion lishes slowly when seeded, it eventually fills the of the spike. They are mostly one to six flowered, interspaces between larger bunchgrasses, thus in- sometimes all fertile, but more commonly with the hibiting the invasion of cheatgrass. Once established, lowermost flower sterile and glumelike. Awns of the Canbar is a vigorous understory species with an abun- glumes and lemmas are 0.8 to 4.7 inches (20 to 120 dance of basal leaves. It makes excellent early spring mm) long, varying from green to red or often purplish. growth, and it spreads well from seed. When planted Glumes are 1.4 to 3.4 inches (35 to 85 mm) long, with other native species it is compatible and not subulate, and extend into slender, spreading, rough- overly competitive. ened awns. Lemmas are 0.3 to 0.4 inch (8 to 10 mm) ‘High Plains’ Selected Germplasm is a composite of long, hairless to strongly short haired, and faintly materials from Montana and Wyoming (Englert and three to five nerved. The main nerve extends into a others 2002). The variety grows to 2 ft (0.6 m) in slender to stout, flexuous awn that may be straight or height. It was selected for use in native seedings to divergent, while two of the lateral nerves extend into provide wildlife habitat and restore native communi- bristles that are about 0.4 inch (10 mm) long (Arnow ties. It is recommended for use on adapted sites in 1987, 1993; Cronquist and others 1977; Hitchcock Montana and Wyoming. 1950; Hitchcock and others 1969). Plants are self- ‘Service’ originated near Whitehorse, Yukon Terri- pollinating and allotetraploid (2n = 28). tory, Canada (Alderson and Sharp 1994). It has not Squirreltail is a member of the Tribe Triticeae, and been seeded in the Intermountain West. It has, how- hybridizes with members of the Elymus, Hordeum, ever, performed well in Alaska when seeded for ero- and Pseudoroegneria complexes. It also produces ster- sion control and revegetation of disturbances on dry, ile hybrids with Great Basin wildrye (Jones 1998). gravelly, and rocky soils. Barkworth and others (1983) included the species in

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Elymus elymoides. Wilson (1963) and Jones and Larson (in press) describe squirreltail as a complex of five taxa. Most widespread is E. e. var. elymoides. E. e. var. californicus is native to the eastern Sierra Nevadas. E. e. var. brevifolius are larger plants, while E. e. var. hordeoides are small and less common. Big squirreltail occurs in the Northwest and southward into Califor- nia and parts of Nevada. The taxa exhibit considerable differences in morphological and phenological characteristics (Hironaka and Tisdale 1972; Jones and Larson, in press) and occupy differing habitats.

Ecological Relationships and Distribution Squirreltail ranges from British Columbia to Saskatchewan and south throughout the Western and Figure 47—Squirreltail is competitive with invasive Central United States and into Mexico. It grows in annual grasses, including cheatgrass and medusahead (RMRS photo). plant communities ranging from salt desert shrub to alpine meadows on sites receiving 8 inches (20 mm)or more of annual precipitation (Horton 1989) at elevations ranging from 2,000 to 11,500 ft (610 to 3,500 m) (Kearney and others 1960). It is a common understory species in sagebrush steppe communities (Jones 1998). (Bradley and others 1992a,b). Squirreltail’s ability to Squirreltail commonly grows on dry, gravelly soils compete with cheatgrass may be favored by fire. that may be saline or alkaline, but some populations also grow on fine-textured soils (Jensen and others Plant Culture 1990; Morris and others 1950). Squirreltail may be an early successional species in overgrazed or otherwise Squirreltail seeds mature from June to September disturbed sites. It is also a mid to late successional depending on site conditions (Link 1993). Mature species (Young and others 1972). Jones (1998) sug- seeds are dispersed following disarticulation of the gests its seral status may be related to both genotype rachis, thus timing of harvest is critical. Doescher and site conditions. (2001) found seed could be harvested when awns are Bottlebrush squirreltail is competitive with the an- divergent and straw colored or about 1 week prior to nual grasses cheatgrass and medusahead and has disarticulation of the rachis. Most viable seed falls been observed to persist on ranges invaded by these near the plant, but fruit morphology, particularly the weeds (Arredondo and others 1998; Beckstead 1994; long awns, permits wind dispersal, and some seeds can Hironaka 1994) (fig. 47). The ability to germinate and be carried as much as 130 ft (40 m) (Beckstead 1994; produce roots at low temperatures permits it to estab- Hironaka and Tisdale 1963; Marlett and Anderson lish in annual grass-infested areas (Hironaka 1994; 1986). The long awns, short seedstalks, and large Hironaka and Sindelar 1975; Young and Evans 1977). amounts of chaffy material make seed collection diffi- Facultative fall or spring germination, early spring cult. Seed fields and extensive wildland stands in growth, extensive root development, and early seed accessible areas are harvested by combining or by first development contribute to the competitive ability of windrowing. Wildland stands are also hand harvested. the species (Reynolds and Fraley 1989). Plants are Awns may be removed with a brush machine, and drought tolerant and winter hardy. trash can be removed with an air-screen machine. Squirreltail is fairly tolerant of burning (Britton and There are about 191,555 seeds per pound (422,302 per others 1990) due to its small size, limited leafy mate- kg) (Plummer and others 1968). Seed can be stored for rial, and coarse, solid stems. The low herbage volume 3 or 4 years (Link 1993). generally limits fire damage to the root crown (Wright Seed can be germinated at constant or alternating 1971; Wright and Klemmedson 1965). Plants are more temperatures of 59, 50/59, or 50/68 ∞F (15, 10/15, or 10/ susceptible to damage when surrounded by accumula- 20 ∞C) without prechilling (Young and Evans 1977). A tions of litter and dead material or when fire frequen- requirement for afterripening may vary among seed cies increase (Wright and Klemmedson 1965). Dam- lots and range from 0 to 4 months (Allen and others aged plants regenerate from surviving portions of the 1994; Beckstead and others 1993; Young and Evans root crown and from on- and offsite seed sources 1977).

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Squirreltail seed may be direct seeded alone or in seed sources a priority for revegetating Intermountain mixtures at depths of 0.25 to 0.5 inch (0.6 to 1.2 cm). rangelands (Hardegree 1994). Squirreltail provides a Seed fields are short lived. Weed control is difficult persistent ground cover and can be used to control and plants are subject to rust. However, seed can be erosion. Direct seeded squirreltail survived for 30 harvested the first year (Archibald and Feigner 1995), years on a big sagebrush/bluebunch wheatgrass site in and production is generally good (Hironaka and Tisdale south-central Idaho (Monsen and Anderson 1993). 1963; Young and Evans 1977). Stand management should emphasize protection from Because of the great variability within this species excessive grazing and protection during heading so complex, seed sources for revegetation efforts must be adequate seed supplies will be produced and the stand selected carefully. Seed should be planted in fall in the maintained (USDA Forest Service 1937; Sanders 1994). Intermountain and Rocky Mountain areas and in late winter to early spring in the Southern Great Plains. Varieties and Ecotypes Plummer and others (1968) ranked germination of squirreltail seedings high, but they considered initial Accessions 9040187 and 9040189 originated in Colo- establishment, growth rate, final establishment, and rado and are being advanced toward release by the persistence of squirreltail seedings only moderate. Upper Colorado Environmental Plant Center in Squirreltail has long been considered one of the Meeker, CO (Alderson and Sharp 1994; UCEPC 2002). more competitive native grasses on cheatgrass range- They were selected for their potential for revegetation lands. Its ability to germinate rapidly across a wide of mined lands and other disturbed rangeland sites. temperature range is considered a factor in its ability Their forage quality is fair to moderate for livestock to establish on disturbed sites (Young and Evans and wildlife, especially in winter. 1977). Allen and others (1994) found that squirreltail Fish Creek, a Sitanion hystrix var. hystrix collection, from a low-elevation site (4,100 ft [1,250 m]) was more was recommended for release as a Selected Germplasm likely to germinate in fall than cheatgrass. In Nevada, by the Utah Agricultural Experiment Station in 2002. Tipton (1994) found that mature squirreltail plants It was collected from a site near Carey, Blaine County, usually initiated growth before cheatgrass. Jones (1998) ID, at an elevation of 4,756 ft (1,400 m) that receives described squirreltail as a potential species for “as- about 15 inches (380 mm) of annual precipitation sisted succession,” a species capable of replacing inva- (Jones and others 2002c). Fish Creek was selected for sive species following wildfire. He listed squirreltail its low proximal awn mass, a trait that may reduce attributes suited to this role, including facultative fall damage during awn removal. In addition, its spikes germination, rapid growth and maturation, root sys- disarticulate at the base, facilitating seed harvesting. tems capable of mining nitrogen, early spring matura- The intended area of use for Fish Creek is the upper tion in areas that dry rapidly, and efficient nitrate Snake River Plain. assimilation. Toe Jam Creek was collected from a site at 6,000 ft (1,829 m) west of Tuscarora, Elko County, NV (Jones Uses and Management and others 2002b). Tuscaraora receives 12.3 inches (312 mm) of annual precipitation. Toe Jam Creek was Squirreltail greens up early in spring and provides developed for use in the northern Great Basin and fair amounts of forage where it grows in dense stands. lower Snake River Plain. It was selected for its large The bristlelike awns can injure grazing animals. Con- seed size and low proximal awn mass. The latter trait sequently, animals avoid these plants from seed ripen- may reduce damage during awn removal, a problem ing until seed dispersal. Late summer herbage and encountered in processing Sand Hollow. Toe Jam especially fall regrowth following rainfall are palat- Creek was identified as Sitanion hystrix var. cali- able and used by livestock and wildlife. Squirreltail is fornicus. The Utah Agricultural Experiment Station a valuable plant for providing energy in winter, as recommended it for release as a Selected Germplasm some leaves remain green (Jensen and others 2001; in 2002. Stubbendieck and others 1992; Vallentine 1971). It is Sand Hollow (Sitanion jubatum or Elymus multi- particularly valuable to big game and small mammals setus) big squirreltail Selected Germplasm originated in winter (Anderson and Shumar 1986; Beale and in Gem County, ID, on a loamy, coarse sand site at an Smith 1970; Kufeld and others 1973). Palatability and elevation of 2,690 ft (820 m) that receives 11 inches use by livestock and big game vary geographically (300 mm) of annual precipitation (Jones and others (USDA Forest Service 1937). 1998b). It is adapted to sandy soils in southern Idaho Squirreltail has considerable potential for reveg- and adjacent regions in Oregon, Nevada, and Utah. etating degraded rangelands, particularly those in- The variety was developed for revegetation of dis- vaded by annual weeds. The USDI Bureau of Land turbed sites and rangelands dominated by exotic Management considers development of squirreltail annuals.

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Sporobolus airoides It grows on dry to moist sites in saline bottoms and in areas with shallow water tables. It tolerates soils with Alkali Sacaton ______up to 3 percent total salts, as well as soils high in Synonyms carbonate and bicarbonate (Blaisdell and Holmgren 1984; Cook and others 1954; Roundy and others 1983; Agrostis airoides Vallentine 1961). Although it can endure alkali, it is Sporoblus wrightii not restricted to alkaline soils. It does poorly on sandy soils, fair on silty soils, and good on clayey soils Description (Vallentine 1971). Plants exhibit moderate fire tolerance, but little shade tolerance. Alkali sacaton is a long-lived, sodforming, native perennial bunchgrass that produces dense clumps Plant Culture that die out in the center over time. In the Southwest this species acts as a warm-season grass, while in the Alkali sacaton reproduces from seeds and tillers. Intermountain area it responds more like a cool- Plants produce numerous seeds from late summer to season grass. Plants are stout and tufted forming October. These may remain viable for many years large clumps clothed with slick, shiny, cream-colored (Blaisdell and Holmgren 1984; Hitchcock 1950). Seeds sheaths at the base. Culms are 16 to 47 inches (4 to 12 must undergo an afterripening period of up to 9 months dm) tall, hairless, round, and usually hollow, but sometimes loosely pith filled. Leaf sheaths are smooth and hairless to slightly silky haired at the upper edges of the margins. Leaf blades are flat to strongly involute, 0.08 to 0.16 inch (2 to 4 mm) wide, smooth below, finely roughened above, and stiff haired near the throat. Ligules are very short with a dense band of hairs. Panicles are 4.7 to 15.7 inches (12 to 40 cm) long, 0.16 to 0.80 inch (4 to 20 mm) wide, open and pyramidal, and often fully exerted or sometimes remaining partially enclosed in the sheath (fig. 48). Spikelets are borne mostly toward the tips of the branches. Glumes are often deciduous. They are tapered to a point, one nerved, roughened or often translucent throughout, and glabrous. The first glume is 0.03 to 0.08 inch (0.7 to 2 mm) long, and the second is about 0.06 to 0.12 inch (1.5 to 3 mm) long. Lemmas are 0.7 to 0.12 inch (1.8 to 3 mm) long and taper to a point; they are usually longer than the second glume (Arnow 1987; Hitchcock 1950; Hitchcock and others 1969; Stubbendieck and others 1986; USDA Forest Service 1937).

Ecological Relationships and Distribution Alkali sacaton occurs throughout the Western United States, the Great Plains, and Northern Mexico. It is most abundant in saltgrass, salt desert shrub, desert shrub, blackbrush, sagebrush-grass, and pinyon-juniper communities. It can also be found in ponderosa pine, mountain brush, semiarid grasslands, and on prairies. It inhabits lower, slightly moist, alkaline flats, where it frequently develops almost pure stands. Stands are also found scattered along drainages in desert and semidesert areas. Throughout the northern portion of its range, it is scattered and less important, but in the Southwest, it occurs in sufficient abundance to be a major forage species. Figure 48—Alkali sacaton is a long-lived bunchgrass Alkali sacaton generally grows in areas receiving 12 that forms large, dense clumps and produces wide, to 14 inches (300 to 350 mm) of annual precipitation. pyramidal panicles (Hitchcock 1950).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 413 Chapter 18 Grasses to relieve dormancy. Local flooding will move seeds establish well and persist in lower elevation shrublands across flood plains to saturated sediments where they that support few species. In these situations it is germinate when favorable conditions are encountered important for controlling weed invasions and protect- (Aldon 1981). ing unstable, erosive soils. This species could become Seeds are small with an average of 1.75 million per much more important for the restoration of arid sites. pound (3.85 million per kg). Purity can be high (98+ percent). Germination of 80 percent is acceptable for Varieties and Ecotypes seed purchases. Seed viability remains high for more than 10 years when seeds are stored dry in a Two varieties of alkali sacaton have been selected warehouse. and released (Alderson and Sharp 1994). Salado was Seed can be broadcast or drill seeded. Extensive selected for use in conservation and range seedings. seedbed preparation is not required. Seeds should be Collected near Clauch, NM, at 5,800 ft elevation planted about 0.25 inch (6 mm) deep. Fall seeding is (1,770 m) it is native to a site that receives 11 to 18 recommended to permit afterripening and release of inches (280 to 460 mm) of annual precipitation. Salado dormancy. Within the same seed lot, some seeds ger- is seeded on valley bottoms, alkaline flats, rocky soil, minate within a few days under favorable condtions, and open plains in the Intermountain West to provide but others are dormant and may not germinate for forage and soil stabilization. It is used on disturbed weeks. Seedling establishment is, therefore, erratic sites where moisture accumulates, such as road right- without some type of irrigation, flooding, or runoff. of-ways, depressions, and gravel pits. Fall seeding is Soils that remain moist for 5 to 10 days to support preferred because germination may be erratic. Seed- germination can produce fair stands (Aldon 1975). ling vigor is generally good. Difficulty in establishment is probably the factor that Saltalk, which originated near Erik, OK, has not most limits the wider use of alkali sacaton. Although been seeded to any great extent in the Intermountain some plantings produce erratic stands, this species West. It is seeded in the Southwest on disturbances in does establish with a good degree of success in quite salt-bearing soils around oil wells. arid communities. It can be seeded in situations where few other species exist. Planting in mixtures with Sporobolus cryptandrus other species, including some shrubs, does not prevent its establishment. Sand Dropseed ______Synonyms Uses and Management Vilfa cryptandra Alkali sacaton provides valuable forage for livestock Agrostis cryptanda in arid and semiarid situations where it can produce a large quantity of forage. As it matures and dries it can Description become coarse and somewhat unpalatable. Cattle and horses do, however, use it in winter months. Sheep will Sand dropseed is a tufted drought-tolerant, use it when forced during drought and on overgrazed nonrhizomatous native bunchgrass that sometimes ranges (Blaisdell and Holmgren 1984; Cook and oth- functions as an annual. It performs as a warm-season ers 1954; Vallentine 1961). Palatability has been rated grass in the Southwest and as a cool-season grass in as good for cattle, good to fair for sheep and horses, and the Intermountain West and other northern areas. It fair to poor for antelope, elk, and deer (Dittberner and can be recognized by the large, distinctive, inflated Olson 1983). sheathes that subtend and partially enclose the inflo- Alkali sacaton is commonly seeded to stabilize dis- rescence. When the sheaths wither, the panicle ex- turbed lands, such as raw mine spoils in the semiarid pands and the seeds are dispersed (fig. 49). Culms are Southwest (Hassell 1982; Oaks 1982; Thornburg 1982). 1.3 to 3.0 ft (4 to 9 dm) tall, erect or prostrate at the It is also planted in riparian zones in a number of base, hairless, and solid. Leaf sheaths are strongly major plant communities in the Intermountain area overlapping, ciliate at the summit, and sometimes (Monsen 1983). This species can be superior to west- hairy along the upper margins. The hairs are 0.04 to ern wheatgrass for seeding in drier portions of the 0.16 inch (1 to 4 mm) long and sometimes occur across southern and northern desert shrub types (Alden the collar. Blades are usually flat with roughened 1981; Plummer 1977). margins and become involute toward the tip. They are Alkali sacaton has been used in range and reclama- 0.06 to 0.20 inch (1.5 to 5 mm) wide and 2 to 6 inches tion seedings in blackbrush, salt desert shrub, sage- (5 to 15 cm) long. Ligules consist of a ring of dense hairs brush, pinyon-juniper, and mountain brush types. It is about 0.02 inch (0.5 mm) long. Panicles are open, 4 to seeded along roadways and riparian corridors. It can 12 inches (1 to 3 dm) long, and 0.6 to 2.4 inches (1.5 to

414 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses

(USDA Forest Service 1937). It is distributed from Ontario, Canada, to Alberta and Washington, and south to North Carolina, Indiana, southern Califor- nia, and northern Mexico (Hitchcock 1950). In the Intermountain West it is most abundant in deserts or lowlands, and it is especially common on sandy soils. This species can be a major component of sagebrush- bunchgrass types, particularly in Idaho and Utah. It is also important in juniper-pinyon, mountain brush, salt desert shrub, blackbrush, chaparral, and ponderosa pine communities and short-grass prairies. Sand dropseed will invade degenerated wheatgrass, fescue, and bluegrass communities. It is able to persist with heavy ungulate use because it has a protected root crown, matures late, and is not a highly preferred species (Johnson and Simon 1987). Sand dropseed responds rapidly to grazing removal, becoming a major component of overgrazed communities (Blaisdell and Holmgren 1984; Cook and others 1954). It can also be one of the first grasses to regenerate on abandoned fields, disturbed sites, waste places, and along roadsides.

Plant Culture Sand dropseed is a prolific seed producer. It spreads well from natural and artificial seeding, and it is often one of the first perennial grasses to reestablish follow- Figure 49—Inflated sheaths that subtend and par- ing drought. Seeds will lie dormant in the soil for many tially enclose the inflorescence of sand dropseed years until favorable conditions for germination and wither when mature, permitting inflorescence ex- establishment occur (USDA Forest Service 1937; pansion and seed dispersal (photo courtesy of Weaver and Albertson 1956). Seedling development is Kevin Jensen, USDA ARS, Logan, UT. rapid, with tillers appearing a few weeks after germination (Weaver and Albertson 1956). Seeds are small with 5.6 million per pound (12.3 million per kg). Seed is often marketed with 98+ 6 cm) wide, with the lower part at least partially percent purity and total germination of 90 to 95 per- enclosed in a sheath. Panicle branches are hairless, cent. However, some seed lots may contain as much as spikelet bearing almost to the base, and usually rather 50 percent hard seed. Seed can be stored for 20 years stiffly ascending to reflexed. Spikelets are cleistoga- and still be 75 percent germinable (USDA Forest mous, or self-fertilized, pale olive green, and hairless. Service 1937). Normally, the older the seed, the better Glumes are tapered to a tip, one nerved, and deciduous the germination and establishment. The species is at maturity. The first is 0.03 to 0.04 inch (0.7 to 1 mm) widely distributed, and ecotypes differ in germination long and awl shaped. The second is 0.06 to 0.09 inch requirements. (1.4 to 2.2 mm) long and broader than the first. Lem- Sand dropseed requires minimal seedbed prepara- mas are about as long as the second glume, 0.06 to 0.10 tion. It can be successfully seeded by drilling or broad- inch (1.5 to 2.5 mm) long, and also tapered to a tip casting and light harrowing on a variety of sites, (Arnow 1987; Cronquist and others 1977; Hitchcock including open exposed soils and unstable distur- 1950; Hitchcock and others 1969; USDA Forest Service bances. Seed should not be planted deeper than about 1937). Sand dropseed roots spread up to 2 ft (0.6 m) 0.24 inch (6 mm), especially in sandy soils. The seed horizontally and 3 ft (0.9 m) vertically (Weaver and coat of sand dropseed is very hard and requires over- Albertson 1956). winter prechilling for germination. Fall plantings can provide natural prechilling and increase the probabil- Ecological Relationships and Distribution ity of spring germination. Seed dormancy can be a deterrent to the establishment of uniform seeded Sand dropseed is an important grass in the South- stands. The growth rate of sand dropseed is considered west, the Intermountain West, and parts of the Snake, good on sandy and salty soils and fair on clayey soils Salmon, and Clearwater drainages in Idaho and Oregon (Stubbendieck and others 1986). It is one of the few

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 415 Chapter 18 Grasses grasses that are well adapted to stabilizing sandy included in revegetation programs wherever it occurs sites. Once plants establish, they are long lived and naturally. It will tend to dominate in some areas such hardy. as sandy outcrops. Sand dropseed is compatible with many other species and can be seeded in mixtures in disturbed salt Varieties and Ecotypes desert shrub, blackbrush, big sagebrush, pinyon-juniper, mountain brush, chaparral, and short-grass prai- There are no released varieties. There is, however, rie communities. In the Southwest, it does well in considerable variation within the species. Care should be areas that are too hot and dry for wheatgrasses. Sand taken to insure that seed of adapted sources is seeded. dropseed is also frequently planted alone. Sand dropseed will establish on sites that receive Stipa columbiana less than 9 inches (230 mm) of annual precipitation. The survival of new plantings of sand dropseed is Subalpine Needlegrass, Columbia dependent upon the availability of spring and early Needlegrass ______summer soil moisture. Plantings that receive good spring precipitation emerge, grow quickly, and be- Synonyms come vigorous and capable of persisting during subse- Achnatherum nelsonii quent periods of summer drought. Stipa viridula var. minor Stipa nelsonii Uses and Management Stipa occidentalis Sand dropseed is fairly productive and palatable to all classes of livestock and many wildlife species, Description especially from spring to midsummer (Daubenmire Subalpine needlegrass is an erect, long-lived, cool- 1970; Hitchcock 1950; Stubbendieck and others 1986; season, native perennial bunchgrass. It produces fine, Vallentine 1961). Its palatability is rated from fair to densely bunched culms that are usually 12 to 24 inches good depending upon associated plants. In many places, (30 to 60 cm) tall, but can be as much as 39 inches (1 this species has been removed due to continued exces- m) tall. Leaf sheaths are strongly ribbed. Leaf blades sive grazing. On ranges in poor to fair condition, sand can be 4 to 10 inches (10 to 25 cm) long and 0.04 to 0.12 dropseed may still provide a major proportion of avail- inch (1 to 3 mm) wide. Leaf edges are flat against the able forage (Alzerraca-Angelo and others 1998; stem and then rolled inward toward the midrib, espe- Stubbendieck and others 1986). It is also used in early cially on younger leaves; they are hairless or slightly summer when plants are succulent. Cattle use it in fall roughened to densely hairy and often strongly stri- following rainfall. This pattern of use favors sand ated. Ligules are short, 0.01 to 0.07 inch long (0.2 to 2 dropseed growth (Johnson and Simon 1987). Plants mm), square, rather firm, and usually longest on the will respond to fall rains and provide succulent forage side. Panicles are narrow, dense, and 2.8 to 8 inches (7 into the winter months (Arnow 1987; Stubbendieck to 20 cm) long (fig. 50). Panicle branches are short and and others 1986). In addition, the herbage cures well remain close to each other. Spikelets produce only one and can provide fair to good winter forage (USDA floret about 0.8 to 1 inch (2 to 2.5 cm) long. The awn is Forest Service 1937). short with the lower portion twisted. The basal end of Wild turkeys feed on sand dropseed in south-central the awn is sharp pointed (Arnow 1987; Hitchcock South Dakota in the Missouri River Breaks region. 1950; Hitchcock and others 1969; USDA Forest Ser- This grass is an important component in the summer vice 1937). diet of Rocky Mountain bighorn sheep, and it is important to deer and elk during most seasons (Rominger Ecological Relationships and Distribution and others 1988). Sand dropseed is an important species that can be Subalpine needlegrass grows in all of the Western used to slow the spread of cheatgrass. Remnant stands States, particularly the Central Rocky Mountains. It and established plants are competitive and can reduce occurs mostly above 8,000 ft (2,440 m), often in dense weed invasion. The species can often be seeded suc- stands, on dry soils of open side hills. It also grows in cessfully on weedy sites if some initial control mea- mountain parks, subalpine herblands, and openings sures are used. Once established, it becomes highly in aspen and conifer forests. It may occur as scattered competitive. It has been useful when planted on big individuals in mountain brush and upper elevation sagebrush and mixed pinyon-juniper sites in central sagebrush areas. There are two recognized varieties: and southern Utah where some cool-season bunch- S. c. var. columbiana occurs from the Yukon to South grasses are less well adapted. This species should be Dakota and south to Texas and California; S. c. var.

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dormancy exist among and within separate wildland collections. Afterripening of dry seeds does occur, and most seeds maintained in storage for 6 months will germinate. Seeds stored for 1 to 7 years usually germinate better than fresh seeds. Cleaned seed lots can usually be planted with most conventional seeders without damaging the seed. Seeds are long and narrow and must be able to pass through openings and regulators that can accommodate long seeds. Seeds must be placed in the soil by drilling or other methods that bury the seed at least 0.25 to 0.50 inch (6 to 12 mm) deep. Broadcasting seeds on the soil surface without some means to cover them is not recommended. Seeds may be dormant, but fall seeding allows seeds to overwinter in the soil, which breaks dormancy and facilitates spring germination. Most species within this genus establish slowly the first season. Although first-year seedlings are normally small, survival may be quite high. Young plants can be slow to reach maturity, especially if seeded in mixtures with other, more rapidly developing species. The Figure 50—Panicles of subalpine needlegrass are slow-developing plants furnish needed ground cover, narrow, compact, and up to 8 inches (20 cm) long; and are less competitive with establishing shrubs and spikelets produce a single floret (Hitchcock 1950). slower developing herbs. Once established, this perennial can persist with a number of other species in rather harsh environments. It is able to exist in the seedling stage with some degree of shade from mixed nelsonii occurs from Alberta to Washington and south overstory shrubs. to Arizona and Colorado (Hitchcock 1950). Subalpine needlegrass often increases on ranges Uses and Management where wheatgrasses and bluegrasses have been removed by excessive use. It resprouts quickly after fires Subalpine needlegrass provides valuable forage for and is able to persist, while other species are often many species of wildlife and livestock. It begins growth slower to respond. It recovers readily with rest from in early spring and remains green throughout a long grazing, often more rapidly than most associated growing season, sometimes until early winter. Most grasses. It will, however, suffer from use during peri- needlegrasses cure well and are grazed during fall and ods of drought. winter (USDA Forest Service 1937). The pointed callus Subalpine needlegrass typically occurs as scattered of subalpine needlegrass sometimes works into the individuals in sagebrush and mountain-brush com- mouth and ears of livestock (Stubbendieck and others munities, but denser stands may be found in subalpine 1992); this causes animals to often avoid the mature grasslands (Vallentine 1961). It often exists on open seed. south and west slopes in association with spreading Palatability of subalpine needlegrass is rated as rabbitbrush, western yarrow, and bluegrasses. Under good for cattle and horses, fair to good for sheep and these conditions it furnishes important ground cover deer, and fair for antelope. It is especially valuable for and soil protection. This needlegrass is slightly to sheep in the spring and for livestock in the summer. moderately damaged by fire (Wright and others 1979). Deer use it in summer, and many other wildlife species It has moderate tolerance to salinity and will grow on consume it throughout the growing season (Vallentine soils ranging from sandy loam to clay loam to dry, 1961). It is particularly valuable for deer and elk in rocky, infertile soils. late fall and winter, as some green remains on the plant. Like most needlegrasses, subalpine needlegrass Plant Culture is palatable early in the season, becoming less so as the foliage becomes coarse and wiry (USDA Forest Ser- Awns can be removed without damage to the seed, vice 1937). This species also provides cover for small and seeds can be cleaned to purities exceeding 90 mammals and birds (Dittberner and Olson 1983). percent. Seed lots contain between 150,000 to 200,000 Subalpine needlegrass is characterized by a slow to seeds per pound (330,000 to 440,000 seeds per kg). moderate seedling growth rate, moderate salinity tol- Considerable variation in seed weights, size, and erance, and the ability to grow on a number of soil

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 417 Chapter 18 Grasses types. It is an important species for revegetation of a appressed silky hairs, and often hairless toward the variety of plant communities that are often seeded to tip. The swollen node (callus) at the base of the floret intermediate wheatgrass and smooth brome. It estab- is very sharp, about 0.12 inch (3 mm) long. The first lishes well and can be used to reestablish native joint in the awn is distinct; the second is rather communities, especially where a mixture of species indistinct and more flexuous. The lower segment is and plant types are desired. tightly twisted and roughened; the terminal segment is not (Arnow 1987; Cronquist and others 1977; Varieties and Ecotypes Hitchcock 1950; Hitchcock and others 1969; USDA Forest Service 1937) (fig. 51). A number of accessions are undergoing development. Accession 9040137 from San Lewis Valley and Ecological Relationships and Distribution Accession 9024804 from western Colorado, are being advanced toward release by the Upper Colorado Envi- Needle-and-thread is widely distributed throughout ronmental Plant Center, in Meeker, CO. These are the Western States and Great Plains. It commonly adapted to areas at elevations of 5,000 to 12,000 ft occurs in sagebrush-grass, pinyon-juniper, and ponde- (1,525 to 3,660 m) receiving 15 inches (380 mm) or rosa pine communities of the Rocky Mountains, Inter- more of annual precipitation. They have potential for mountain West, and semidesert plains and foothills of soil stabilization, rangeland seedings, and revegeta- the Southwest. Needle-and-thread is most often found tion of disturbed sites in the West (Alderson and Sharp on well-drained soils. 1994). With overgrazing, needle-and-thread is one of the first species to disappear from native pastures and Stipa comata Needle-and-Thread ______Synonyms Hesperostipa comata Stipa juncea Stipa tweedyi

Description Needle-and-thread is an erect, long-lived native perennial cool-season bunchgrass. It is named for the exceptionally long, twisted, and tapering awns that resemble threaded sewing needles. Plants are tufted with culms that are 1.0 to 3.6 ft (3 to 11 dm) tall and often slightly hairy at the nodes. Sheaths are smooth to roughened, strongly ribbed, and usually longer than the internodes. Leaf blades are involute, or if flat, up to 0.12 inch (3 mm) wide and 4 to 12 inches (10 to 30 cm) long, roughened on the upper surface, and smooth to slightly roughened beneath. Ligules are 0.04 to 0.24 inch (1.1 to 6 mm) long, more or less minutely hairy, and usually tapering to a tip. They may become lacerate with age. Panicles are 5 to 15.5 inches (13 to 40 cm) long, narrow, and usually partly enclosed in an inflated sheath. Panicle branches are usually slender and ascending, sometimes with drooping spikelets. Glumes are long and narrow, tapering to a fine point. They are five nerved, slightly convoluted, hairless, and papery, with translucent margins and tips. The Figure 51—Needle-and-thread is named for the first glume is 0.7 to 1.4 inches (1.8 to 3.5 cm) long, and florets with their extremely long awns that resemble the second is 0.6 to 1.3 inches (1.5 to 3.4 cm) long. threaded sewing needles (photo courtesy of Loren Lemmas are 0.32 to 0.55 inch (8 to 14 mm) long, pale St. John, USDA NRCS, Aberdeen Plant Materials green to yellowish or brownish, sparsely covered with Center, Aberdeen, ID).

418 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses range sites. Likewise, it is one of the first species to some seeds. Even when cleaned, the sharp tip on the respond and reoccupy native communities following seed and awn fragments can cause handling and seed- rest from grazing. ing problems. Mixtures of needle-and-thread and non- awned seed of other species can be planted with most Plant Culture seeders. Needle-and-thread depends on seed for reproduc- Uses and Management tion. In favorable areas, it reproduces well and it is able to withstand grazing in spring and fall if plants Needle-and-thread is an important range grass for are allowed to mature seed in summer. On drier livestock and wildlife species because of its early ranges, however, it does not reproduce readily and it is spring and late fall greenup, palatability, volume of easily killed by overuse. forage produced, and widespread distribution. The Needle-and-thread can be established in sagebrush forage value of needle-and-thread varies depending on and some salt desert shrublands. It is drought tolerant the region, season, and associated plants. Its palat- and requires well-drained soil. Seeds can be drilled or ability is rated very high in early spring before the broadcast, but they must be covered with at least 0.25 awns develop and again after the seeds are dispersed. inch (6 mm) of soil. Needle-and-thread can be seeded It is especially valuable because it begins development successfully with most native perennial grasses and in early spring when other grasses have not yet initi- broadleaf herbs. It should not be seeded with species ated growth. An abundance of herbage is produced and that are vigorous competitors. Seedling development will remain green through summer and fall if ad- of needle-and-thread can be slow, but even during low- equate moisture is available. Awns can become a precipitation years some seedlings will generally es- problem when the fruit matures because the sharp, tablish. It may fail to develop dense or dominant barbed lemmas can injure grazing animals. New growth stands for the first 5 to 10 years after planting, even is produced in summer and fall if rainstorms occur. when seeded alone. After this time, the stands develop Needle-and-thread cures well and is often utilized as slowly, even during years of low annual rainfall. As the winter forage. This is one of the most important stand becomes denser, persistent perennials may be grasses of semiarid and arid shrublands of the Inter- suppressed. Seeded stands of needle-and thread have mountain region. Its loss due to grazing has resulted displaced intermediate wheatgrass on some pinyon- in an increase of annual weeds, particularly cheatgrass. juniper planting sites in central Utah approximately Recovery of this perennial is critical to control weeds. 25 years after the initial seeding. This species has also This species is an important element of the shrub- recovered naturally on sites where pinyon and juniper bunch grass communities and pinyon-juniper wood- trees have been removed and the native understory lands, and reestablishment of functional communities has been allowed to recover. Heavy grazing by live- requires inclusion of this species. stock or big game can delay or suppress natural recovery. Varieties and Ecotypes Needle-and-thread has not prevented shrub seedling establishment when seeded with big sagebrush, There are no released varieties. Care should be antelope bitterbrush, Stansbury cliffrose, or curlleaf taken to ensure that seed of adapted populations is mountain mahogany. It can be utilized to establish a planted. Great Plains collections do not do especially mixed array of native species without necessitating well in the Intermountain West. separate planting operations. Stands of this species provide sufficient competition to restrict the spread of Stipa lettermanii annual weeds. It recovers quickly after burning and does not allow weeds to enter after fires. Letterman Needlegrass ______Needle-and-thread seed is marketed with the awn Synonyms removed. There is an average of 115,000 seeds per pound (253,000 seeds per kg). Some seed dormancy Stipa viridula var. lettermanii exists, but seed with 75 to 85 percent germination and 80 to 85 percent purity is generally marketed. Seed Description can be stored for a number of years with little loss of viability. The long, tough awns limit commercial pro- Letterman needlegrass is an erect, fine-stemmed, duction and use of this species. Some growth forms densely-tufted, native perennial bunchgrass that of- produce seeds with flexible awns that do not break ten forms large clumps. Stems are slender and 8 to 20 from the seeds. Growth forms with brittle awns can be inches (20 to 50 cm) tall. Leaves are threadlike and cleaned and seeded with much less difficulty. Tech- somewhat twisted. Culms are 10 to 33 inches (2.5 to niques and equipment have been developed to de-awn 8.5 dm) tall. Culms and leaf sheaths are hairless or

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 419 Chapter 18 Grasses minutely roughened. Leaf blades are involute, rounded, Plant Culture rarely flat, and up to 0.08 inch (2 mm) wide and 4.0 to 8.0 inches (10 to 20 cm) long, stiff haired above and Seeds may be drilled or broadcast seeded, but they hairless to minutely roughened below. Ligules are must be covered with about 0.25 inch (0.60 cm) of soil. 0.01 to 0.08 inch (0.2 to 2 mm) long and rounded to Fall seeding is recommended because seed dormancy square. Panicles are narrow and about 2.8 to 9.5 can be overcome by overwinter stratification. inches (7 to 24 cm) long with erect branches and Letterman needlegrass has been seeded success- spikelets. Glumes are subequal, 0.20 to 0.41 inch (5 to fully on disturbed sites to control erosion, revegetate 10.5 mm) long, and taper gradually to a tip. They are mines, and restore lost stands. As is the case for other often purple with translucent margins, hairless or needlegrass species, seedlings develop slowly. sometimes minutely roughened, and three nerved. Letterman needlegrass can be planted with most na- Lemmas are only slightly hardened, 0.20 to 0.26 inch tive species that are not highly competitive. Letterman (4.5 to 6.7 mm) long, and pale with soft, slender, needlegrass usually produces seeds each year, and it somewhat appressed hairs that become longer toward will spread into unoccupied sites. the tip. The callus often has dense, white hairs less than 0.04 inch (1 mm) in length. Awns are 0.40 to 0.87 Uses and Management inch (1.0 to 2.2 cm) long, slender, and twice bent with Letterman needlegrass is an important component the lower segment loosely twisted and minutely rough- of many sagebrush, mountain brush, and aspen com- ened; the terminal segment is 0.3 to 0.5 inch (7 to 12 munities where it may vary in density and abundance. mm) long and hairless. The basal end of the seed It recovers well after fires and can persist with some tapers to a pointed tip (Arnow 1987; Cronquist and shade or competition. Although it is rated as only fair others 1977; Hitchcock 1950; Hitchcock and others for cattle and poor for sheep, it is an important forage 1969; USDA Forest Service 1937). species on many sites (USDA Forest Service 1937). Letterman needlegrass is often misidentified as Heavy grazing by cattle or sheep can cause a decrease, subalpine needlegrass because the two plants are or sometimes a short-term increase, in density. similar in size. They can, however, be differentiated by Letterman needlegrass provides valuable forage for the width and length of the palea in relation to the wildlife and domestic livestock. Plants begin growth length of the lemma. The palea of Letterman early in the year, and they remain green throughout needlegrass is about two-thirds the length of the the summer and late fall. This enables Letterman lemma, and it is often exposed. The palea of subalpine needlegrass to be utilized when many other grasses needlegrass is less than half as long as the lemma. The are less available or palatable. Most needlegrasses, distribution of the two species is about the same, both including Letterman needlegrass, cure well and can be grow in all Western States, with the Central Rocky utilized in fall and winter. This species is particularly Mountains as their center of distribution. Letterman needlegrass is, however, less abundant at the northern and southern ends of its range, and it does not extend to altitudes as high as does subalpine needlegrass.

Ecological Relationships and Distribution Letterman needlegrass is found on dry soils in open mountain parks, mountain meadows, subalpine grasslands, and in openings in aspen and conifer forest from Oregon and Montana, south to Arizona and New Mexico (fig. 52). It is also found in midelevation pinyon-juniper, sagebrush, and mountain brush communities. It grows best on sandy loam, loam, or clay loam soils 20 inches (51 cm) or more in depth (Dittberner and Olson 1983). It tends to increase under heavy grazing by sheep and decrease under heavy cattle use. Letterman needlegrass is a climax or late seral species in a number of grassland, sagebrush, mountain-shrub, and pinyon-juniper communities. It also grows well in early successional assemblages, and is Figure 52—Letterman needlegrass grows in mountain well represented in many early to late seral plant meadows, forest openings, and subalpine grasslands associations (Schott 1981). (RMRS photo).

420 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses important as late fall and early winter forage for species in grasslands, sagebrush deserts, and pinyon- big game; the presence of some green material within juniper, oak, and ponderosa pine woodlands (fig. 53). the plant attracts these animals and influences their In sagebrush deserts it is most common in communi- distribution. ties receiving 6 to 16 inches (152 to 406 mm) of annual precipitation (USDA Natural Resources Conservation Varieties and Ecotypes Service 2003). It is fairly widespread in the northwestern portion of the Great Basin. There are no releases. Thurber needlegrass is adapted to soils ranging from well-drained clay loams to coarse-textured soils Stipa thurberiana that are fairly neutral (USDA Forest Service 1937; USDA Natural Resources Conservation Service 2003). Thurber Needlegrass ______Plants are tolerant of drought and cold, but they are Synonyms not salt or shade tolerant (USDA Natural Resources Conservation Service 2003). Achnatherum thurberianum Response of Thurber needlegrass to fire is variable, Stipa occidentalis but it is generally less resistant to burning than other bunchgrasses (Wright and others 1979). Burning is Description more damaging early in the season than later (Britton and others 1990; Ganskopp 1988; Uresk and others Thurber needlegrass is a densely tufted perennial with culms 1.2 to 2.6 ft (3.5 to 8 dm) tall that are minutely hairy, at least at the nodes. Leaf sheaths are striated and usually hairless, but sometimes minutely haired below. Blades are needlelike, involute, minutely roughened, and 3.9 to 9.8 inches (10 to 25 cm) long. Ligules are 0.08 to 0.28 inch (2 to 7 mm) long, thin and translucent, hairless to sparsely haired, and tapering at the tip. Panicles are narrow and about 2.8 to 9.5 inches (7 to 24 cm) long with erect branches that are relatively few flowered. Glumes are subequal and 0.31 to 0.63 inch (8 to 16 mm) long with the second slightly shorter than the first. They are three- to five- nerved, often purplish and translucent above, and taper to a soft tip. Lemmas are 0.24 to 0.34 inch (6.2 to 8.7 mm) long and sparsely covered with silky, appressed hairs. The callus is about 0.04 inch (1 mm) long, with dense, appressed hairs. Awns are 1.2 to 1.8 inches (30 to 45 mm) long and bent twice. The lower segments are twisted with stiff, feather-like hairs up to 0.08 inch (2 mm) long. The terminal segment is 0.6 to 1.2 inches (16 to 30 mm) long and roughened to hairless. Thurber needlegrass flowers from late May through July. It is known to hybridize with Oryzopsis hymenoides, producing the hybrid Stipa bloomeri (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969; Johnson 1945).

Ecological Relationships and Distribution Thurber needlegrass ranges from eastern Washing- ton, Oregon, and California, to northern Nevada, southern Idaho, southwestern Montana, and northeastern Wyoming (Arnow 1987, 1993; Cronquist and others Figure 53—Thurber needlegrass occurs as a 1977; Hitchcock 1950; Hitchcock and others 1969). It dominant or common associated species on dry may occur as a dominant or associated mid to late seral big sagebrush sites (RMRS photo).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 421 Chapter 18 Grasses

1980). The presence of dry and sometimes abundant sagebrush/Thurber needlegrass cover type in eastern leafy culms contributes to surface heating and burn- Oregon prior to nesting (Barnett and Crawford 1994). ing of the root crown during wildfires (Wright and Thurber needlegrass is valuable for cattle and sheep Klemmedson 1965). Accumulations of dead vegetative in early spring when other forage is not available. material contribute to fire damage at all seasons. Palatability and nutritive value of the species is great- Plants may be killed or living tissue may survive near est early in the season, declining as plants and seeds the edges of the root crown. Plants then regenerate mature and the forage becomes coarse (USDA Forest from tillers if postfire weather conditions permit Service 1937). Leaves are used to a greater degree (Britton and others 1990; Koniak 1985; Uresk and than stems. The species tends to be avoided as the others 1976, 1980). Surviving plants recover slowly seeds mature because the awns can irritate or injure and exhibit reduced vigor and productivity for up to 3 the ears, eyes, and noses of grazing animals. Because years. The number of inflorescences and seed produc- of this, considerable seed generally matures, even in tion, however, may be enhanced by burning (Uresk areas grazed by livestock (Jensen and others 2001; and others 1980), contributing to postburn regenera- USDA Forest Service 1937). Thurber needlegrass re- tion from seed (Hironaka and others 1983). mains green after other grasses dry and provides Thurber needlegrass seedlings compete poorly with useful forage in fall. those of more competitive annual and perennial grasses Thurber needlegrass stands are damaged by exces- (Evans and Young 1978; Uresk and others 1980; Young sive grazing and fire. Overgrazing in spring can be and Evans 1978). Dependence on residual or offsite particularly detrimental because defoliation, particu- seed sources and low germination and seedling vigor larly during the boot stage, can weaken plants tend to slow recovery following burns. Consequently, (Ganskopp 1988). The species tends to increase with Thurber needlegrass often does not become abundant protection from grazing (Robertson 1971). Thurber until communities are mid to late successional (Koniak needlegrass can be used for reclaiming mine sites and 1985). other disturbances on dry sites where it is adapted. Such uses, however, are limited by a lack of seed. Plant Culture Varieties and Ecotypes Thurber needlegrass ripens in mid to late summer (Sampson 1924). Seed production is generally low. There There are no releases. are about 225,000 seeds per pound (496,035 per kg) (USDA Natural Resources Conservation Service 2003). Haferkamp and McSwain (1951) found that germina- Stipa viridula Trin. tion was enhanced by incubation at a fluctuating (68/ Green Needlegrass ______86 ∞F) (20/30 ∞C) temperature (16 hrs/8 hrs) with exposure to light during the high-temperature periods. Synonyms Thurber needlegrass can be seeded on adapted sites Nassella viridula receiving as little as 10 inches (254 mm) of annual Stipa parviflora precipitation (Thornburg 1982). Litter has been found Stipa nuttalliana to reduce germination and early growth of Thurber Stipa sparta needlegrass seedlings, but it was also associated with improved growth of establishing seedlings 4 weeks Description after germination (Schlatterer and Tisdale 1969). Seed- ling vigor is low to moderate (Hironaka and others Green needlegrass is a long-lived, native, cool-sea- 1983; USDA Natural Resources Conservation Service son, perennial bunchgrass that is tolerant of drought, 2003), and spread of stands from seed is slow. cold winters, and a wide range of soil types. It is a tufted perennial with culms 1.6 to 3.6 ft (5 to 11 dm) Uses and Management tall. Leaf sheaths are persistent and hairless or hairy along the margins. Blades are flat to involute, smooth Thurber needlegrass provides valuable forage for live- or roughened, and 0.80 to 0.24 inch (2 to 6 mm) wide. stock and wildlife. Deer, wild horses, pronghorn, black- Ligules are 0.02 to 0.12 inch (0.5 to 3 mm) long. Many tailed jackrabbits, and other species graze the herbage plants are functionally pistillate and bear only small, (Fagerstone and others 1980; McInnis and Vavra 1987; sterile anthers; thus, the species may be semi-dioe- Trainer and others 1983; Uresk and others 1976). Birds cious. Panicles are narrowly oblong, 4 to 10 inches (1 to and other small animals use its seeds. Thurber 2.5 dm) long, and approximately 0.8 inch (2 cm) wide. needlegrass and associated species provide cover for Panicle branches are erect or only slightly divergent. many small vertebrates and invertebrates (Stanton 1974). Glumes are translucent throughout, or nearly so, equal Sage-grouse hens were found to use a Wyoming big or subequal, and 0.32 to 0.51 inch (8 to 13 mm) long,

422 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 18 Grasses tapering to a long, hairlike tip. They are prominently collections exhibit relatively high levels of seed dor- three to five nerved and hairless. The lemma is brown mancy. It is not uncommon to purchase fresh seed and leathery at maturity, 0.20 to 0.26 inch (5 to 6.5 with less than 10 percent germination and 90 percent mm) long, and narrowed toward the tip into a hairless, hard seed. Considerable afterripening occurs during whitish “neck” approximately 0.04 inch (1 mm) long. dry storage with germination percentage increasing The awn is 0.8 to 1.4 inches (2 to 3.5 cm) long and twice up to 7 years after harvest (Rogler 1960). Conse- bent, with the two lower segments twisted and mi- quently, better stands are normally obtained with nutely roughened. The callus is up to 0.04 inch (1 mm) older seed. Fall seeding allows seeds to overwinter in long, with a very short, blunt, hairless tip (Arnow the soil. This can relieve dormancy and allow seeds to 1987; Cronquist and others 1977; Hitchcock 1950; germinate and seedlings to establish in spring. For Hitchcock and others 1969; USDA Forest Service this reason spring seeding is not recommended. Seed 1937). Green needlegrass occasionally hybridizes with can be drilled or broadcast but must be covered be- Indian ricegrass, but the offspring are sterile (Hitchcock cause maximum germination occurs in darkness and others 1969). (Fulbright and others 1982). Seed need not be seeded deeper than about 0.25 inch (6 mm). Ecological Relationships and Distribution Germination potential and seed maturity are correlated. Immature seed, which is light gray to yellowish Green needlegrass is a perennial bunchgrass dis- green, has a lower average weight than does mature tributed from British Columbia to Minnesota, Kansas, seed, which is a uniform gray (Kinch and Wiesner New Mexico, Arizona, Nevada, and western Washing- 1963). Average viability of immature seed is approxi- ton. It occurs more commonly in Montana, Wyoming, mately 22 percent compared with 71 percent for ma- and Colorado than in the other Western States. Green ture seed (Kinch and Wiesner 1963). needlegrass grows on dry slopes, plains, prairies, and Within the Intermountain region, green needle- foothills at lower elevations and often may be found grass has performed well when planted in aspen, growing along roadways. It is also common in moun- mountain brush, and mountain big sagebrush com- tain meadows, open woodlands, and on hillsides at munities. Sites that receive at least 14 inches (350 mm) higher elevations. At the southern edge of its distribu- of annual precipitation will generally support this tion, green needlegrass grows in dry parks and can- grass. Vigor of green needlegrass seedlings is good yons and with ponderosa pine (USDA Forest Service (Kinch and Wiesner 1963). However, when planted 1937). It is a prominent species in many grassland and with pubescent or intermediate wheatgrass, its ini- sagebrush communities, and it grows well on aban- tial performance has been suppressed by these doned agricultural lands (Wasser 1982). sodformers. Green needlegrass exists as an early or late seral Mature green needlegrass plants are very competi- species. It is adapted to a wide range of soils. It grows tive with annual weeds, but new seedlings are not. especially well on clay and moderately alkaline soils When planted in areas with cheatgrass and other derived from calcareous shales. It is somewhat less annuals, the weedy species must be eliminated or adapted to loam and sand, and it is not very tolerant controlled to facilitate the establishment of this peren- of saline conditions (Wesser 1982). This species exhib- nial. New plants will establish in areas where annual its considerable drought tolerance, which is nearly kochia and mustards dominate. New seedlings are equal to that of western wheatgrass. It can be found also able to establish and survive in mixed grass-forb growing in areas with as little as 10 inches (250 mm) seedings. Green needlegrass typically establishes well of annual precipitation. Optimum growth occurs with but can decrease over time; it is often planted in 12 to 20 inches (310 to 510 mm) of precipitation. It mixtures of cool-season, native or introduced grasses grows, however, in areas with as much as 30 inches or forbs that tend to persist. (760 mm) precipitation and in areas with water tables only 1 to 2 ft (30 to 60 cm) deep (Wasser 1982). Green Uses and Management needlegrass is moderately tolerant of flooding, only weakly tolerant of shade, and somewhat tolerant of fire, Many Stipa species produce awns or calluses that depending on the season of burn and site conditions. cause injury to grazers. This is not the case with green needlegrass. Green needlegrass provides valuable for- Plant Culture age for many species of wildlife and all classes of livestock (fig. 54). It is palatable to cattle and horses Green needlegrass flowers and disperses seeds from season long, but is used by sheep primarily in spring July to September. Cleaning seeds to remove the awns and early summer (USDA Forest Service 1937). and other debris can be completed without damage to Green needlegrass begins growth in early spring the seed. There are an average of 181,000 seeds per and remains green until late summer when most other pound (400,000 per kg). Seed of most green needlegrass grasses have dried. In many areas, annual growth

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 423 Chapter 18 Grasses

early-season grazing (Wasser 1982). Wasser (1982) recommends moderate grazing of stands and resting a different unit for 2 months during the growing season each year. When green needlegrass occurs with warm- season grasses, seeded stands should be grazed during the spring and fall or in winter (Wasser 1982). Green needlegrass has been successfully seeded throughout the Intermountain area. It has done especially well on disturbed sites in the mountain brush zone and lower elevation aspen zones. Green needlegrass has been widely used for rehabilitating disturbed sites. This species has a wide range of adaptation and performs well in rangeland plantings in the aspen and mountain brush communities throughout the Intermountain West. It has also performed well on mine and rangeland seedings in eastern Wyo- ming, Montana, and western Colorado. Green needlegrass is particularly useful for erosion control. It has been successfully seeded on abandoned farmlands and pastures. Seedlings develop slowly, and should not be grazed for up to 4 years following seeding. This species can be planted in mixtures to restore native communities. It does not compete to exclude native shrubs and herbs.

Varieties and Ecotypes ‘Cucharas’ was collected near Walsenburg, CO, and is recommended for use in the Central Great Plains (Jones and Larson, in press; Jones and others 2002d, 2004b). It was selected for its high germination without a wet prechill treatment. Cucharas was recommended for release as a Selected Germplasm by the Utah Agricultural Experiment Station in 2002. Figure 54—Most common in the northern Great Plains, ‘Lodorm’ originated near Bismarck, ND (Alderson green needlegrass provides forage for many wildlife and Sharp 1994), and is seeded for forage, soil stabili- species (photo courtesy of Tom Jones, USDA ARS, zation, and revegetation of mined sites and other Logan, UT). disturbances. Lodorm was developed for the Northern Great Plains, but it is also being seeded in the Inter- mountain West. It is adapted to areas receiving 14 or more inches (355 mm) of annual precipitation and begins as early as March (Wasser 1982), with most elevations above 9,000 ft (2,745 m) in mountain brush, growth occurring May and June. Late-season regrowth ponderosa pine, and aspen openings. It is seeded on can be good if fall precipitation is sufficient. This bottomlands, flats, benches, and along streams. Lodorm species is generally found in mixed stands with other is adapted to silt and clay soils, and it is tolerant of grasses, although dense stands are common. As a dense clays. It will also grow on loamy and sandy soils. result, succulent forage is available over a longer This variety exhibits rapid vegetative recovery, high period of time where green needlegrass is part of the palatability, low seed dormancy, and rapid germina- community. Green needlegrass cures well and can be tion. It is compatible with other natives and can be used for fall and winter grazing. sown in mixtures. Fall seeding is recommended to Green needlegrass is moderately resistant to heavy relieve seed dormancy. This grass is very winter hardy, grazing (Wasser 1982), and generally recovers quickly tolerant of fire and short-duration flooding, and weakly (Kinch and Wiesner 1963). It can be harmed by intense shade tolerant.

424 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Richard Stevens Stephen B. Monsen Chapter 19

Forbs for Seeding Range and Wildlife Habitats

Forbs are abundant in all vegetative types throughout the Intermountain West. Most are found intermixed in grasslands and as understory plants in shrub and forest types. Forbs provide ground cover, soil stability, community (flora and fauna) diversity, nutritious forage, and are of aesthetic value. Forbs should be seeded in most range, watershed, mine reclamation, highway, recreational site, restoration and revegetation projects. Many commonly used forbs have been introduced to the Intermountain West and are now under cultivation for seed production. The principal introduced forbs planted on range and wildlands are alfalfa, small burnet, and cicer milkvetch. Lewis flax, Rocky Mountain penstemon, and Palmer penstemon are the major native forbs being commercially grown and planted. Additional species are being commercially grown in seed fields as demand increases. Seeds of many native species are collected from wildland stands but generally are in short supply.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 425 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Chapter Contents by Species

Achillea millefolium ssp. lanulosa (western yarrow) ...... 430 Artemisia ludoviciana (Louisiana sage) ...... 430 Aster chilensis (Pacific aster) ...... 431 Aster glaucodes (blueleaf aster) ...... 433 Astragalus cicer (cicer milkvetch) ...... 433 Balsamorhiza sagittata (arrowleaf balsamroot) ...... 435 Balsamorhiza macrophylla (cutleaf balsamroot) ...... 436 Balsamorhiza hookeri var. neglecta (hairy balsamroot) ...... 436 Coronilla varia (crownvetch) ...... 437 Geranium richardsonii (Richardson geranium) ...... 438 Geranium viscosissimum (sticky geranium) ...... 439 Hedysarum boreale (Utah sweetvetch) ...... 439 Helianthella uniflora (oneflower helianthella) ...... 440 Heracleum lanatum (cow parsnip)...... 441 Ligusticum porteri (Porter ligusticum) ...... 442 Linum perenne spp. lewisii (Lewis flax) ...... 443 Lomatium triternatum (nineleaf lomatium) ...... 444 Lomatium kingii (nuttall lomatium) ...... 445 Lupinus sericeus (silky lupine) ...... 446 Lupinus argenteus (silvery lupine) ...... 447 Medicago sativa (alfalfa) ...... 447 Medicago falcata (sicklepod alfalfa) ...... 447 Melilotus officinalis (yellow sweetclover) ...... 451 Mertensia arizonica (tall bluebell) ...... 452 Onobrychis viciaefolia (sainfoin) ...... 453 Osmorhiza occidentalis (sweetanise) ...... 454 Penstemon palmeri (Palmer penstemon) ...... 454 Penstemon cyananthus (Wasatch penstemon) ...... 456 Penstemon eatonii (Eaton penstemon) ...... 456 Penstemon humilis (low penstemon) ...... 457 Penstemon pachyphyllus (thickleaf penstemon) ...... 457 Penstemon rydbergii (Rydberg penstemon) ...... 457 Penstemon strictus (Rocky Mountain penstemon) ...... 457 Sanguisorba minor (small burnet) ...... 457 Senecio serra (butterweed groundsel) ...... 459 Solidago canadensis (Canada goldenrod)...... 460 Sphaeralcea grossulariifolia (gooseberryleaf globemallow) ...... 461 Sphaeralcea coccinea (scarlet globemallow) ...... 462 Trifolium (clovers) ...... 463 Trifolium fragiferum (strawberry clover) ...... 463 Trifolium hybridum (alsike clover)...... 464 Trifolium pratense (red clover) ...... 464 Trifolium repens (white clover) ...... 464 Viguiera multiflora var. multiflora (Showy goldeneye)...... 465 Viguiera miltiflora var. nevadensis (Nevada goldeneye) ...... 466

Important characteristics of a number of forbs are The following forb species have the potential for listed in table 1. Seeding recommendations for princi- improving range and wildlife habitats in the Inter- pal vegetative types and conditions are discussed in mountain West. A brief description along with infor- chapter 17. Seed characteristics and information mation regarding ecological relationships and distri- about collection, cleaning, and storage are provided bution, cultural requirements, use, and management in chapter 24. Forbs adapted to different vegetative of each species are included. types and conditions are included in the seeding recommendations.

426 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

(con.)

egetative

A,PP,MB,MS

JP,MS,BS,WS

A,PP,MB

SA,A,MB

SS,BB PP,MB,MS PP,MB,JP, WS,SS,BG,BB MS,BS SS,BB PP,MB,JP, MS,BS BS,WS MB,JP,MS,BS BS,WS MS,BS

SA,A,MB

JP,BS

MB,JP,MS,BS

MS,BS,WS

WS,BB

ility adapted

adapt- species is

333

323

-V

433

344

41 444

444

354

514

543

Early Compati- type

Ratings

443

444

423 444

334

432

525

334

433

334

114 334 2444334442423442SA,A,WM 233

434

114

553

335

243

4423445333342434333MB,JP,MS,BS,

5432344443333434323JP,MS,BS,WS,

413 3432444334442423442SA,A,WM 243 4433343224421423552SA,A,WM 3232343254421443442SA,A,WM 433

3422355524542334233A,PP,MB,JP

424

413

514

4 2 2 2 3 2

Ease of Ease of Initial Seedling Final spring Summer Toler- bility with Seed Flood Shade Range which the

handling Ease of trans- Germi- estab- growth estab- Persis- Natural Forage palata- palata- ance to Evergreen- other produc- Soil toler- toler- of

—Characteristics of Selected Forbs.

Iris, German 1 4 5 2 1 4 5 5 4 2 5 4 4 5 3 1 4 4 4 4 SA,A,PP,MB, (common iris) Ligusticum, Porter 4 3 2 4 3 3 3 4 3 3 4 4 4 1 3 4 3 2 3 2 A,PP,MB

Helianthella, oneflower

Goldenrod, low 2 2 4 4 3 4 4 4 4 4 4 4 5 2 3 4 5 3 3 3 A,PP,MB Goldenrod, Parry 2 2 4 4 3 4 5 4 4 4 4 4 5 2 3 4 5 3 3 4 A,PP,MB Globemallow, 3 5 3Groundsel, butterweed 3 3 3 4 4 4 3 3 2 4 2 4 3 4 2 2 3 JP,MS,BS,WS, gooseberryleaf Globemallow, scarlet Balsamroot, hairy 4 4 1 3 1 1 4 5 2 4 5 1 4 1 4 4 3 1 3 2 PP,MB,MS Bluebell, tall 3 3 4 3 2 4 3 4 3 5 5 5 4 1 2 3 4 4 3 2 SA,A,WM Globemallow, stream Bouncing-bet 5 5 5 4 4 4 4 5 5 4 3 3 4 2 3 4 5 4 3 3 PP,MB,MS Burnet, small 5 5 4Fireweed 5 5 1 5 2 3 4 3 5 4 5 3 5 5 3 5 2 3 5 5 4 4 4 5 3 3 5 2 1 2 4 3 4 PP,MB,JP,MS, 4 3 4 3 SA,A,WM,PP, Goldeneye, Nevada 3 3 4 4 4 4 3 3 5 3 4 4 4 1 4 3 3 2 2 3 MS,BS,WS Goldeneye, showy 3 3 4 4 4 4 4 3 5 3 4 4 4 1 4 4 3 4 3 3 SA,A,PP,MB,JP, Balsamroot, cutleaf Cinquefoil, gland 4 4 5Flax, Lewis 4 5 5 4 5 5 4 5 4 5 5 5 4 3 4 3 3 5 5 1 3 5 5 4 3 5 4 4 4 1 4 3 4 A,WM,PP,MB 3 2 3 4 PP,MB,JP,MS, Clover, alsike 5 5 5 4 4 3 4 4 4 4 4 4 4 2 3 3 4 5 3 1 WM Goldenrod, Canada Clover, strawberry 5 5 5Geranium, Richardson 5 4 3 4 5 5 2 4 4 4 2 4 3 4 5 3 2 WM,IS Table 1 Columbine, Colorado Geranium, sticky 2 3 4 3 Cow parsnip, common Giant hyssop, nettleleaf Aster, blueleaf 2 2 5 4 4Crownvetch 4 5 5 5 5 5 5 4 4 4 4 4 5 4 5 5 2 5 2 4 3 4 5 4 4 4 3 2 4 A,WM,PP,MB, 2 4 5 3 4 3 A,PP,MB,MS Daisy, common oxeye Aster, Englemann 2 2 4 3 3Deervetch, birdfoot 3 3 4 4 4 5 5 4 4 4 4 4 5 4 4 3 2 4 4 5 2 5 4 5 4 5 4 2 2 3 A,WM,PP,MB 3 5 3 3 3 A,PP Aster, Pacific 2 2 5 4 3Eriogonum, cushion 3 4 5 5 3 5 5 5 2 3 3 5 4 3 4 A,WM,PP,

Balsamroot, arrowleaf Species seed seeding planting nation lishment rate lishment tence spread yield bility bility grazing ness species tion stability ance ance ab

Alfalfa, range type 5 5 4 5 5 5 4 5 4 5 5 4 4 3 5 5 4 2 4 5 SA,A,PP,MB,JP,

Alfileria 2 4 2 4 4 5 3 2 5 3 5 3 4 1 3 4 3 1 3 3 JP,MS,BS,

Angelica, small leaf

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 427 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

(con.)

Vegetative

PP,MB,MS

PP,MB,JP, MS,BS

JP,MS,BS PP,MB A,PP,MB,JP, MB,JP,MS,BS MS,BS PP,MB

BS,WS MB,JP,MS, BS,WS MS,BS

MB,JP,MS,BS, MS,BS WS PP,MB,JP, MS,BS SA,A,WM, PP,MB BS,WS

BS,WS BS,WS

BS,WS

BS,WS,BB

ility adapted

adapt- species is

443

453

143

344

343

324

434

343

334

434

Early Compati- type

Ratings

433

343

354

342

443

344

553

443

334

233

3322344325431333532SA,A,WM, 343

5233345444443434343PP,MB,JP,MS,

234

5432344424343433333PP,MB,JP,MS,

543

5443244443433334333PP,MB,JP,MS, 543

5454433544445343234PP,MB,JP,MS,

543

4 3

5 5

Ease of Ease of Initial Seedling Final spring Summer Toler- bility with Seed Flood Shade Range which the

handling Ease of trans- Germi- estab- growth estab- Persis- Natural Forage palata- palata- ance to Evergreen- other produc- Soil toler- toler- of

(Con.)

Table 1

Lomatium, nuttall 3 3 3 3 3 3 4 3 3 2 5 2 4 1 3 4 3 2 3 3 A,PP,MB Lupine, Nevada 5 5 2 3 3 3 4 4 3 4 4 4 4 1 4 3 3 2 3 4 JP,MS,BS,WS Lupine, silky 5 5 2 3 3 3 4 4 3 4 4 4 4 1 4 3 3 3 3 4 A,PP,MB,MS Lupine, silvery 5 5 2 3 4 3 4 4 3 4 4 4 4 1 4 3 3 3 3 4 A,PP,MB,MS Medick, black 4 5 5 5 5 4 4 4 4 3 4 4 4Solomon-plume, fat 2 4 5 5 4 3 2 A,WM,PP,MB, Lomatium, nineleaf Milkvetch, cicer 4 5 5 3 3 3 4 5 5 5 4 4 5Sweetanise 3 3 4 3 5 2 5 3 3 4 4 3 4 SA,A,WM,PP, 4 4 4 4 4 4 4 2 4 4 3 4 5 3 SA,A,WM,

Milkvetch, Snake River Sage, Louisiana 2 3 5 4 2 4 4 4 5 3 2 3 5 3 4 4 5 4 3 4 SA,A,WM,PP plains Milkvetch, tall 4 5 2 3 4 4 4 4 4 2 3 4 4 3 3 3 4 4 4 2 PP,MB,JP, Sage, tarragon 2 2 4 3 1 4 4 4 3 3 2 2 4 4 3 3 4 3 3 3 SA,A Species seed seeding planting nation lishment rate lishment tence spreadPainted-cup, yield bility 2 bility grazing 4 ness species 1 tion stability ance 2 ance 2 ab 3 4 3 2Sainfoin, common 3 5 4 5 4 5 2 5 1 4 2 4 2 4 1 3 3 4 3 5 3 5 SA,A,WM,PP, 4 4 3 3 4 3 3 4 3 PP,MB,JP, northwestern Salsify, vegetable-oyster Peavine, flat 4 5 4 3 3 3 3 4 3 4 4 4 3 3 5 2 5 4 4 3 SA,A,PP,MB Peavine, perennial 4 5 4 3 3 4 4 4 4 3 3 5 3 2 3 3 4 4 4 3 SA,A,PP,MB Peavine, thickleaf 4 5 4 3 2Penstemon, toadflax 3 4 5 3 4 4 4 4 2 4 2 5 4 4 4 SA,A,PP,MB Peavine, Utah 4 5 3 3 2 3 4 4 3 4 5 5 3 2 4 2 4 4 4 3 SA,A,PP,MB Penstemon, Eaton 5 5 5Penstemon, sidehill 3 5 5 5Penstemon, Wasatch 4 4 4 4 3 3 5 3 3 4 4 4 4 4 3 2 3 3 3 3 3 4 3 3 3 4 3 3 4 PP,MB,JP,MS, 4 3 3 3 PP,MB,JP,MS,

Penstemon, littlecup Penstemon, thickleaf

Penstemon, low 5 5 5 3 4 3 5 5 5 2 4 4 4 3 4 3 5 3 3 3 PP,MB,JP,MS,

Penstemon, Palmer

Penstemon, Rocky 5 5 4 5 4 3 4 3 4 3 4 4 4 4 4 5 4 3 3 3 A,PP,MB,JP, Mountain Penstemon, Rydberg

428 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Vegetative

SA,A,WM, PP,MB

BS,WS

MB,JP,MS

SA,A,WM, BG,BB PP,MB, JP,MS,BS,WS,

PP,MB

MB,JP,MS,BS

ility adapted

adapt- species is

344

243

244

345

sh; JP = Juniper-pinyon; MS = Mountain big sagebrush; BS = Basin big sagebrush;WS = Wyoming big sagebrush;

443

343

Early Compati- type

Ratings

322

234

334

552

432

535

Ease of Ease of Initial Seedling Final spring Summer Toler- bility with Seed Flood Shade Range which the

handling Ease of trans- Germi- estab- growth estab- Persis- Natural Forage palata- palata- ance to Evergreen- other produc- Soil toler- toler- of

(Con.)

Key to vegetative type: SA = Subalpine; A = Aspen-conifer; WM = Wet and semiwet meadows; PP = Ponderosa pine; MB = Mountain bru Key to ratings: 1 = Poor; 2 = Fair; 3 = Medium; 4 = Good; 5 = Excellent.

Sweetvetch, Utah 2 4 3 3 3 3 5 4 3 4 4 4 4 3 4 4 3 3 3 3 PP,MB,JP,MS,

Valerian, edible 3 3 2 2 2 2 3 5 2 3 4 4 4 1 4 3 3 4 4 3 SA,A,WM,PP,

Vetch, American 4 5 4 4 2 4 4 5 3 4 4 4 4 3 5 4 4 3 4 3 SA,A,WM,PP,

Vetch, bramble 4 5 4 5 3 2 4 5 3 3 4 4 4 3 5 4 4 3 4 3 BS,WS,SS,

Yarrow, western 4 4 5 4 4 4 5 5 5 3 4 4 5 4 4 5 5 5 4 5 SA,A,WM,PP,

Sweetroot, spreading

Table 1 Sweetclover, white 5 5 3 5 5 5 2 2 3 3 3 4 3 3 4 5 2 4 3 4 SA,A,WM,

Sweetclover, yellow

Species seed seeding planting nation lishment rate lishment tence spread yield bility bility grazing ness species tion stability ance ance ab

SS = Shadscale saltbush; BG = Black greasewood; BB = Blackbrush; IS = Inland saltgrass.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 429 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Family Compositae types (USDA Forest Service 1937). Western yarrow has good fire tolerance, some shade tolerance, and is Achillea millefolium ssp. lanulosa__ fairly resistant to drought. It grows in sandy to loam Western Yarrow soils that range from weakly basic to weakly acidic (USDA Forest Service 1937; Wasser 1982). Yarrow In the Intermountain West the majority of the yar- spreads by rhizomes and seed, and is especially useful rows fall into three taxonomic units: Achillea on disturbed or misused sites. Tolerance to grazing is millefolium ssp. millefolium, which was introduced good. from Europe and is widely cultivated in North America, and A. millefolium ssp. lanulosa, a native to North Plant Culture America, which is divided into two varieties—var. lanulosa (western yarrow) that occurs from the valley Western yarrow can be seeded successfully by drill- floor to above timberline and var. alpicola that is ing or broadcasting. Seeds are small (4.1 million per lb generally found above timberline (Welsh and others [9.0 million per kg] at 100 percent purity) and should 1987). All three taxonomic units intergrade. Western not be planted more than 0.25 inch (6 mm) deep. Seeds yarrow, a perennial member of the sunflower family, germinate uniformly, and produce vigorous seedlings. has flowers that are borne in flat-topped cymes (fig. 1) Seed is usually marketed with about 50 percent purity that are white, grayish, and barely pink, all with and 80 percent germination. It is recommended that yellow centers (Hermann 1966). The leaves and flow- seed be stored no more than 2 years, due to loss of ers are aromatic. viability. The species is easily transplanted. Western yarrow can be seeded successfully in mixtures with Ecological Relationships and Distribution other species (Stevens and others 1985c). Western yarrow is found in Southwestern Canada, Uses and Management in all States west of North Dakota, south to Northern Mexico, and from the low plains to subalpine commu- Western yarrow has fair forage value for sheep, nities. Greatest areas of occurrence are in mountain somewhat less for cattle, and is used by small and brush, ponderosa pine, aspen, and open timber large game animals (Hermann 1966; Kufeld 1973; Kufeld and others 1973; USDA Forest Service 1937; Wasser 1982). It provides important habitat for upland game birds. This species has good potential for revegetation and for stabilization of disturbed sites from big sagebrush through the subalpine zones. West- ern yarrow establishes and spreads well in sites occupied by annual weeds. It persists with heavy grazing, and often recovers well by natural seeding. Areas seeded to western yarrow should not be grazed for at least 1 to 2 years following seeding.

Varieties and Ecotypes There is considerable variation, and locally adapted ecotypes should be planted.

Family Compositae Artemisia ludoviciana ______Louisiana Sage (wormwood) There are a number of subspecies and numerous varieties and ecotypes of Louisiana sage in the Inter- mountain West (Holmgren and Reveal 1966; Welsh and others 1987). Considerable differences occur among the various subspecies, varieties, and ecotypes. Loui- siana sage is an herbaceous native perennial that is aromatic and very rhizomatous, with stems that grow from 1 to 3 ft (30 to 91 cm) tall. Leaves are Figure 1—Western yarrow in a mixed mainly cauline, entire, lobed or pinnately incised, and grass forb community. borne on the stem.

430 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Ecological Relationships and Distribution Louisiana sage (fig. 2) occurs throughout North America in all vegetation types from sagebrush through alpine and tundra zones. This species does well on shallow as well as on deep, slightly acidic to basic soils. It prefers well-drained soil, but also grows in riparian communities and on other moist sites. Louisiana sage does well on disturbed exposed soils as well as on sites supporting some cover. As a pioneer species, it creates microenvironments suitable for the invasion of other species.

Plant Culture Flowering occurs in mid to late summer, depending Figure 2—Louisiana sage. on location, with seed maturing in late August to early November. Seed can be collected by hand stripping, beating, or combining. Most native stands do not produce an abundance of seeds on a regular basis. extensive use or trampling from grazing animals. Seed is cleaned by debearding and screening. There Stand density fluctuates, particularly at higher eleva- are about 2.5 million seeds per lb (5.5 million per kg) tions where the species may be suppressed by over- at 100 percent purity. Seed is generally marketed at story shading. It recovers well after fires and can be 10 to 15 percent purity and 80 percent germination. used to prevent the entry of weeds by providing ground Seed viability declines rapidly with more than 3 to cover after a disturbance. 4 years of storage. Seed can be drill seeded or broadcast seeded, fol- Varieties and Ecotypes lowed by very shallow soil coverage. Drill seeding requires a purity of greater than 40 percent for seed to There is one variety. ‘Summit’ was selected for its flow through most seeding units. Seed can be broad- its ability (1) to establish from sprig planting and cast with as low as 10 percent purity, depending on the direct seeding, (2) increase or spread rapidly by root- type of broadcaster used. Seed that is fall broadcast on ing, (3) provide quick cover and soil stabilization, (4) disturbed soil generally does not require mechanical grow on harsh sites, and (5) create microenvironments coverage because natural soil sluff is sufficient to bury suitable for the invasion of other species (Stranathan the seed. Louisiana sage can be seeded as a single and Monsen 1986). ‘Summit’ is particularly useful as species or as a component of a mixture. It also estab- a ground cover or erosion control species for sites lishes easily from sprigs. above 5,000 ft (1,500 m) elevation.

Uses and Management Family Compositae Louisiana sage plays an important roll in providing Aster chilensis______soil cover and stabilization in areas where it is adapted (Hermann 1966; McCulloch 1973). It can be an early Pacific Aster invader on disturbed sites including mine spoils Pacific aster is a native perennial of the compositae (Monsen 1975; Monsen and Plummer 1978). Where it family. It is described by Harrington (1964) as devel- is seeded or becomes established, it acts as a nursery oping short to long root stocks and stems that are crop that allows establishment of other species. Ellison usually less than 2.5 ft (0.8 m) tall, with long, linear (1951) considered it an important invader of over- pubescent leaves. Few to numerous heads occur with grazed sheep range and as a soil stabilizer on the blue, violet, or rarely white flowers. Seeds or achenes Wasatch Plateau in central Utah. It can be used to are usually hairy with many white pappus bristles. stabilize riparian disturbances including dry mead- Plants bloom from July to September. This is an ows and areas subjected to infrequent flooding. extremely diverse species. Welsh and others (1987) Livestock, small and large game, and rodents use consider Pacific aster a generalized taxon with no this forb (Hermann 1966; Kufeld and others 1973; well-defined diagnostic features. Hitchcock and oth- McCulloch 1973). It has considerable tolerance to ers (1955) recognized three subspecies: spp. hallii, grazing and trampling; however, when occurring on which occurs in the Oregon Cascades, and in Washing- shallow soils or disturbed sites, it should not receive ton west to the coast, spp. chilensis, which is found

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 431 Chapter 19 Forbs for Seeding Range and Wildlife Habitats throughout the Cascades and Sierras; and spp. adscendens, which occurs in the Great Basin and eastern Oregon, and on the Snake River plains.

Ecological Relationships and Distribution Pacific aster (fig. 3) is a native perennial that can spread rapidly by seed or rhizomes. This forb occurs throughout the Intermountain region at elevations between 5,000 and 9,500 ft (1,500 to 2,700 m). It occupies areas from the sagebrush foothills to the subalpine zone (Harrington 1964). It is most commonly found on dry, open sites in the sagebrush-grass and pinyon-juniper types.

Plant Culture Figure 3—Pacific aster in full bloom. Pacific aster can be established by direct seeding, sprigging, and with nursery or field-grown transplants (Stevens and others 1985c). Seed is best planted small plants do not attract heavy use and provide by broadcasting; however, if it is cleaned to at least excellent ground cover. 50 percent purity, it can be drilled. Picker or thimble seeders can be used to plant the uncleaned, trashy Uses and Management seed. Special care must be taken to ensure that seeds are not planted more than 0.25 inch (6 mm) deep. To Pacific aster is one of the first forbs to “green up” in ensure that viable seed is planted, seed should not be the spring, causing it to be highly sought by livestock stored more than 3 years. Harvested seed usually and big game. Game and livestock do, however, make consists of the achene and attached pappus. The some use of this species during all seasons because the pappus is removed by gentle hammermilling, or plants retain some green material most of the year with a debearder. Seed is usually not cleaned to (Hermann 1966; Kufeld and others 1973; Plummer more than 50 percent purity, and most seed is sold and others 1968). Hermann (1966) and USDA Forest at 10 to 14 percent purity. At 100 percent purity, a Service (1937) reported that Engelmann aster is more pound of seed consists of just over 2.5 million seeds palatable than Pacific aster to most animals. Our (5.5 million per kg). observations indicate similar differences, but recognize Seeds of Pacific aster ripen throughout the late that Pacific aster receives moderate to heavy sea- summer and fall. Not all seeds ripen uniformly; conse- sonal grazing. quently, considerable debris and immature seeds are Pacific aster is one of the most widely distributed often collected. Seeds can be separated from the debris native forbs within the Intermountain region. Selec- with special cleaning equipment. Seed should be pur- tions obtained from the low foothill sagebrush commu- chased based on the percent of viable seed. nities are well adapted to semiarid situations. Pacific Seeds of Pacific aster germinate readily. Nearly all aster is one of a few forbs that can be easily seeded viable seeds germinate within 15 days. Seedlings within the arid region occupied by big sagebrush. It appear early. They are able to compete with most persists well, furnishes good midsummer forage, and seeded herbs and persist under adverse conditions. can be used to control cheatgrass. It recovers well from Young plants usually do not attain large stature the wildfires and spreads naturally. first growing season, but they do form an extensive To date, Pacific aster has normally been seeded as a minor component of most seed mixtures, usually at root system. 0.25 lb seed per acre (0.3 kg per ha). However, it can be Although the seeds are small, they establish well seeded at higher rates of 2 to 5 lb per acre (2.2 to 5.6 kg with only minimum seedbed preparation if they are per ha) to provide more summer forage. Pacific aster seeded with the pappus attached. Seeding with the does not restrict seedling establishment of associated pappus on, however, is very difficult. herbs, and its own seedlings are not impaired by the Neither livestock grazing nor rodent and insect use presence of other seeded species. Consequently, in- have been observed to damage new seedings. New creasing the seeding rate of this plant usually results plantings are not heavily used by rabbits and can be in a corresponding increase in plant density. established where rabbit populations are high. The The strong rhizomes and root system enables the plant can also be seeded on unstable slopes because species to be very useful in stabilization of disturbed

432 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats areas. Pacific aster is a useful species for planting exposed sites with shallow soils, particularly midwinter game ranges. It can be successfully seeded on areas that are difficult to treat with mechanical equipment. It is well suited to infertile, rocky soils and sites subject to erosion. Pacific aster is a useful pioneer species and nurse crop. The plant is able to occupy harsh sites and improve surface stability, enhancing seedbed conditions for other species. New seedlings are able to establish amid the seemingly dense ground cover provided by this forb. It is not unusual to observe seedlings of other plants establishing and surviving within a clump of Pacific aster. Consequently, the plant is used to furnish cover on steep slopes where Figure 4—Blueleaf aster at seed maturity. This pinyon and juniper have been chained. The plant is stand was established by broadcast seeding on a also valuable for seeding or transplanting roadways, raw roadfill in the mountain brush type. mines, and other major disturbances. Areas seeded to Pacific aster should not be grazed for at least two growing seasons following planting. Creeping rhizomes enable this species to withstand considerable grazing and trampling once plants are established. Pacific aster has only been sparingly used in controlled grazing studies, but it is apparent that this species can be managed to provide midsummer and winter forage. In addition, the plant can be seeded to control cheatgrass and to furnish needed cover on critical semiarid ranges.

Varieties and Ecotypes There are no releases.

Family Compositae Aster glaucodes ______Figure 5—Blueleaf aster stabilizing a severely Blueleaf Aster eroded cut. Blueleaf aster (fig. 4), a native perennial, has about the same forage value and seed characteristics as Pacific aster. However, its seeds are more than three times as large (500,000 per lb [1.1 million per kg]), and Family Fabaceae the species is more widely distribted, commonly occur- Astragalus cicer ______ring in all vegetative types from salt desert shrublands and saline seeps to spruce-fir forests. It furnishes an Cicer Milkvetch open spreading clump, with short seed stalks and wide leaves. It is well adapted to harsh sites (fig. 5) and A native of Eurasia, cicer milkvetch is a perennial tends to exist alone. This species is better adapted to member of the pea or legume family. It has vigorous calcareous soils and saline conditions than Pacific creeping rhizomes and a short taproot. The leafy, aster. When seeded with principal forage grasses, it succulent, decumbent stems are fairly large in diam- does not provide a dominant cover. It is grazed by big eter. They grow to about 5.5 ft (1.7m) in length, and game and livestock, but it is not heavily used. Like originate from crown and rhizome buds. Flowers are Pacific aster it persists amid heavy infestation of born in compact heads and are pale yellow to white. grasshoppers when other succulent herbs are usually Blooming occurs in midsummer, and seeds mature in consumed. If seed of both species were more available, late fall. The inflated, bladder-shaped, leathery black they would be recommended for planting range, wa- pods contain several seeds. Seed pods persist on the tershed, and wildlife habitats. plant into the winter.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 433 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Ecological Relationships and Distribution Seed should be inoculated with appropriate inoculant before seeding (Carleton and others 1971; Hafenrichter Cicer milkvetch (fig. 6) is an introduced, long-lived and others 1968), and should not be planted more than perennial from northern Europe. It is bloat free and 0.25 inch (6.4 mm) deep. Because of its hard seedcoat does not accumulate toxic alkaloids (Davis 1982a; and dormancy, seeding should be done in the fall. Rumbaugh 1984; Rumbaugh and Townsend 1985; Scarification can improve seeding success. However, USDA Soil Conservation Service 1968, 1972a; Williams once seeds are scarified they will not retain viability and others 1976). It is safe for grazing (Rumbaugh over extended periods. 1983). Seed of cicer milkvetch has been stored for up to 14 This species has vigorous spreading rhizomes, good years without any loss of viability (USDA Soil Conser- frost and drought tolerance, and produces consider- vation Service 1968a). This species is easily trans- able succulent forage with a high protein content and planted. Cicer milkvetch has done well seeded in a low crude fiber level (Davis 1982a). It grows in mixtures and has been successfully interseeded into slightly acidic and basic soils, and does especially well various native and introduced grass communities in soils derived from limestone. This forb establishes (Rumbaugh and others 1981; Stevens and others and grows on thin, infertile soils and on disturbed sites. 1981b). Because of its lower seedling vigor (Rumbaugh Spring growth starts about 2 weeks later than alfalfa 1984), cicer milkvetch does best when seeded in well and continues about 2 weeks longer in the fall. Foliage prepared seedbeds that are relatively free of competi- is green and succulent throughout the summer and tion. This species does not reach full productivity fall months. Basal leaves remain green through win- until the third year following seeding. Once estab- ter. This species has more frost tolerance than alfalfa lished, cicer milkvetch is very competitive and long- and remarkable resistance to insects and disease. lived. It has been demonstrated that cicer milkvetch Cicer milkvetch requires at least 13 inches (33 cm) can stimulate and increase forage production of of annual precipitation, and does especially well with associated species (Johnson and others 1983). 16 inches (40 cm) or more. It is tolerant of water tables within 3 ft (91 cm) of the surface and does fairly well Uses and Management on wet sites. It has some shade tolerance, and is well adapted to sagebrush-grass and pinyon-juniper sites Seed of cicer milkvetch is produced commercially with sufficient precipitation. It also does well in the and is generally available. Acceptable purity and ger- mountain brush type (Plummer and others 1955; mination for commercial seed is 95 and 85 percent. Rumbaugh and Townsend 1985) and in openings in Cicer milkvetch is palatable to livestock and big game, aspen and subalpine areas. and seeds are eaten by small birds, deer, rabbits, sage- grouse, and pheasants (Plummer and others 1968; Plant Culture USDA Soil Conservation Service 1968, 1972a; Wasser 1982). Once established, this species has excellent Cicer milkvetch can be broadcast or drilled seeded. grazing tolerance. This species can be classified as a There are about 113,000 seeds per lb (249,000 per kg). semievergreen because basal leaves are green all year. The rhizomatous characteristics of cicer milkvetch, along with its semiprostrate nature and profuse flowering, make it ideal for use around summer homes, on disturbed sites, in campgrounds and parks, and in areas with high aesthetic value. This species is useful as a forage and conservation plant, providing excellent ground cover and stabilizing eroding sites. It can be seeded for intensive pasture management. The plant has considerable value for seeding semiwet sites that are used by upland game birds for summer forage. It also provides winter and nesting cover for upland game birds.

Varieties and Ecotypes ‘Lutana’ was released in 1970 and has the character- Figure 6—Cicer milkvetch growing in associa- istics described for the species. ‘Monarch’ was selected tion with smooth brome, Gambel oak, and for its superior seedling emergence. ‘Oxley’ is a 1971 mountain big sagebrush. Canadian release (Rumbaugh and Townsend 1985).

434 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Family Compositae Balsamorhiza sagittata ______Arrowleaf Balsamroot Arrowleaf balsamroot (fig. 7) is a broadleaf perennial with a deep, woody taproot. It is the most abundant balsamroot in the Intermountain area. Simple deltoid or sagittate leaves are entirely gray-green and heavily pubescent. Flowering stems produce a few reduced leaves. Flowers or heads are usually borne solitary on extended seed stalks. Both disk and ray flowers are produced, but they lack a pappus (Welsh and others 1987). Flowering occurs in early April, and seed matures in late May and early June (Harrington Figure 7—Seed increase planting of arrowleaf 1964). Leaves dry up between late June and the end of balsamroot. July.

Ecological Relationships and Distribution A native perennial forb, arrowleaf balsamroot can be found growing on well-drained silty to loamy soils Seeds lack a fluffy or bristly pappus common to in sagebrush, mountain brush, and ponderosa pine many other compositae. Seeds are long, smooth, and communities, and open slopes in aspen and conifer- easily cleaned. Wildland stands are often infested ous forests. This forb is usually found at elevations with seed damaging insects, and entire seed crops can between 1,000 to 9,600 ft (305 to 2,900 m) (Harrington be damaged. Seed should be carefully inspected prior 1964; Wasser 1982). It occurs in solid stands or mixed to collection. Plummer and others (1968) suggest treat- communities. Arrowleaf balsamroot is the most wide- ing seed production areas with insecticides to reduce spread species of balsamroot. It ranges from the Sier- seed predation However, treatment must be timed to ras east to Colorado and north to southern Canada on avoid elimination of pollinators. Seed in storage should acidic and alkaline soils (Hitchcock and others 1955; be treated to prevent insect damage. Seed stored Plummer 1977). It dominates many extensive foothill properly in an open warehouse retained good viability areas, often growing as nearly pure stands (USDA after 5 years (Stevens and Jorgensen 1994). Forest Service 1937). Mature seed will remain on the plant for a short Arrowleaf balsamroot is intolerant of shallow water time, but is readily eaten by rodents (Everett and tables, but withstands brief periods of soil saturation. others 1978b). Immature and insect-damaged seed It is strongly drought resistant, winter hardy, and will persist on the plant for longer periods. Too often, tolerant of semishade, open sunlight, and of grazing the persisting poor-quality seed is mistakenly har- (Wasser 1982). vested and marketed. Immature and damaged seed may often be normal size. However, it is light weight Plant Culture and can be separated from fully developed seed by gravity, wind separation, or flotation seed cleaning Arrowleaf balsamroot is one of the first plants to techniques. Good quality lots will have 95 percent initiate growth in the early spring. Plants flower in purity and at least 40 percent germination. April and usually attain maximum stature in late Arrowleaf balsamroot seeds can be drill seeded quite April or May, and seeds ripen in May and early June. easily with conventional equipment. Seeds should not Seed heads are formed individually on extended soli- be placed more than 0.33 to 0.5 inch (8 to 13 mm) deep. tary peduncles, and if plants are not heavily grazed, Seeds can be broadcast planted but only if they are large quantities of seed can be produced. Seeds ripen covered. Because of seed dormancy and rodent prefer- uniformly and are large, with just over 55,000 per lb ence for this species, late fall seeding is recommended (120,000 per kg). Seed is easily harvested by hand (Eddleman 1978; Everett and others 1978b; McDonough stripping or beating. Extensive areas of level terrain 1976; Young and Evans 1979). Arrowleaf balsamroot and dense stands can be harvested with a combine or seeds are often planted alone or with slower develop- reel-mounted harvester (Plummer and others 1968; ing herbs and shrubs. This species should not be Stevens and others 1985c). Seed can also be produced seeded directly with more aggressive developing under cultivation (fig. 7). herbs, but should be planted in separate rows or

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 435 Chapter 19 Forbs for Seeding Range and Wildlife Habitats broadcast onto sites free of competition. If planted Like most other natives, arrowleaf balsamroot with grasses or other broadleaf herbs, arrowleaf grows on many different soil types and in many pre- balsamroot plants may survive, but would require cipitation zones. Consequently, various ecotypes likely 10 years or longer to reach maturity. occur. Considerable differences are noted among popu- Although seedlings germinate and emerge quickly, lations in regard to palatability, seed production, and young plants develop slowly. Even under ideal con- drought tolerance. Seeds are normally produced each ditions thick, dense stands usually do not develop year from upper elevation sites where precipitation is from artificial seedings. Causes for low return are more reliable; however, seeds should be acquired from not fully known, but insects and rodents can cause sites similar to the area proposed for treatment. The considerable damage. Seeded plants may produce genus also appears to lack genetic barriers to hybrid- only two to three small leaves for a number of years ization, and intergradation occurs when any two taxa following planting. Plants are slow to increase in exist together (Welsh and others 1987). stature, but they are extremely hardy and persistent.

Uses and Management Family Compositae Balsamorhiza macrophylla______Arrowleaf balsamroot is an important forage plant for deer, elk, cattle, and sheep, especially on spring Cutleaf Balsamroot ranges where it greens up early (Harniss and Wright 1982; Hermann 1966; Holecheck and others 1982; Cutleaf balsamroot has large leaves 12 to 24 inches Kufeld 1973; Kufeld and others 1973; Mueggler and (30 to 61 cm) long that are divided into broad, entire, Stewart 1980; Plummer and others 1968). The plant or few-toothed segments 2 to 5 inches (5 to 13 cm) long is also grazed later in the season with considerable (Welsh and others 1987). The leaves have long, soft use made of the heads during and following flower- hairs and a somewhat unpleasant odor. ing. In many plant communities, a major portion of Cutleaf balsamroot generally occurs at slightly total forage production is arrowleaf balsamroot. higher elevations and in more moist conditions than Plantings of arrowleaf balsamroot do well when arrowleaf balsamroot. It is not distributed as widely seeded within areas of its natural occurrence. At- as arrowleaf balsamroot and is less palatable to graz- tempts to extend or seed this species on areas outside ing animals. Cutleaf balsamroot ranges from northern of its natural range or on mine or roadway distur- Utah and southeastern Idaho to western Wyoming bances have been unsuccessful. This forb is encoun- and southwestern Montana (Hitchcock and others tered in low, semiarid conditions with Wyoming big 1955). Seed collection, cleaning, and storage as de- sagebrush, but has been difficult to seed in these scribed for arrowleaf balsamroot. Seeds are somewhat circumstances, usually because of using unadapted larger than those of arrowleaf balsamroot with ecotypes, heavy grazing, and competitive effects of about 33,000 per lb (73,000 per kg). Seed can only be associated species. The most successful seedings have stored for 3 years without significant loss of viability occurred in the lower elevation mountains in sage- (Stevens and Jorgensen 1994). Cutleaf balsamroot is brush and mountain brush communities. better adapted to areas of slightly higher elevations Arrowleaf balsamroot can be used to improve forage and more moist conditions than arrowleaf balsamroot. conditions on game and livestock spring and summer Neither species should be seeded in place of the other. ranges. It undoubtedly has been eliminated from some Site differences are not easy to differentiate. situations, but is extremely hardy and persistent even under heavy use. Because of a deep fleshy taproot, arrowleaf balsam- Family Compositae root can withstand drought, trampling, fire and con- Balsamorhiza hookeri var. siderable grazing pressure. Natural reproduction will neglecta ______occur if seeds are allowed to mature. Seeded areas should not be grazed for at least two growing seasons Hairy Balsamroot following planting. New plants are slow to mature, requiring 3 to 4 years to flower on the most preferred Compared to arrowleaf balsamroot, hairy balsam- sites, and 7 to 8 or even 10 years on the lower root has smaller leaves 4 to 12 inches (10 to 31 cm) long precipitation sites. that are divided into narrow segments 0.5 to 2 inches (1 to 5 cm) long (Welsh and others 1987). Varieties and Ecotypes These segments are often divided a second time into even smaller segments. The leaves are covered with There are no releases. coarse, short, stiff hairs.

436 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

This species has a more limited distribution than arrowleaf or cutleaf balsamroot, occurring from eastern Oregon to southwestern Nevada (Hitchcock and others 1955). Welsh and others (1987) report two varieties of Hooker balsamroot are found in Utah, these occur in separate regions, but grow on a variety of sites. Seed collection, cleaning, and storage are as described for arrowleaf balsamroot. Seed production is much lower than for arrowleaf or cutleaf balsamroot. In addition, plants usually do not exist in large dense patches, but are scattered and intermixed with other herbs. Plants usually grow as an understory with shrubs, and have been useful in seeding pinyon-juniper and mountain brush sites. Sources that grow in drier circumstances offer opportunity to select and develop materials adapted to sagebrush and salt desert Figure 8—Crownvetch stabilizing a road cut. shrublands.

Family Fabaceae Coronilla varia ______Plant Culture Crownvetch Seeds are produced in segmented pods. Cleaned seeds are separated from the pod, and should be Crownvetch (fig. 8) is an introduced perennial le- inoculated with the proper rhizobium prior to seed- gume with strong rhizomes and a deep taproot (Leffel ing (Magness and others 1971; Wasser 1982). There 1973). Varigated white to purple flowers start to blos- are about 135,000 seeds per lb (298,000 per kg) at som in early summer and blooming continues for 4 to 100 percent purity. Acceptable purity and germina- 5 weeks. Seed matures in early fall in pencil-like pods tion for commercial seed is 95 and 75 percent. Seeding that break into segments as they dry. Each segment is best accomplished in the fall on a well-prepared contains one yellow to dark red, rod-shaped seed. seedbed. Stratification requirements can be overcome Flowers are arranged in an umbrella or crown-like with fall seeding. Seed can be drilled or broadcast, but fashion. Basal leaves are green early in the spring and it should not be covered deeper than 0.25 inch (6 mm). remain green into the winter. This forb can be seeded with other species. When seeded with a grass mixture, it is best seeded separately Ecological Relationships and Distribution through the legume seedbox. Excellent establishment can be obtained from transplanting or sprigging. Seed- Crownvetch is a long-lived, cold-tolerant perennial ing at a rate of 2 to 4 lb per acre (2.2 to 4.5 kg per ha) from the Mediterranean region. This species is not a is satisfactory when seeded in a mixture. true vetch because it does not have tendrils for climbing. It does, however, tend to climb if supported by shrubs like mountain big sagebrush, Gambel oak, or Uses and Management serviceberry. This forb does fairly well with 21 inches Crownvetch produces strong, spreading, fleshy rhi- (530 mm) annual precipitation and thrives with over zomes. This extensive system enables crownvetch to 30 inches (760 mm) of precipitation. Crownvetch exhib- be an excellent soil stabilizer and an aggressive its some shade and fire tolerance. It prefers slightly spreader. It has been used to stabilize watershed and acidic soils but has also done well on slightly basic mine disturbances at locations receiving over 16 inches calcareous soils. It does not persist on poorly drained (40 cm) of annual precipitation. The plant is character- soils. Crownvetch can be seeded in mountain brush, pon- ized as having a succulent dark green appearance, derosa pine, and aspen communities. It persists well profuse flowering, and excellent ground cover charac- at lower elevations once established. Although it per- teristics. These features make crownvetch a prime forms best when grown on fertile soils, it demonstrates landscape species for summer home developments, unusual ability to persist, spread, and remain produc- roadways, campgrounds, and areas of high recre- tive when planted on mine disturbances and sites ational use. lacking top soil. It is especially useful for stabilizing Palatable forage is produced by crownvetch for all erosive sites. This species is very competitive. Over time classes of livestock and wildlife (Wasser 1982), with it will dominate an area and exclude other species. little or no bloat hazard (Leffel 1973; USDA Soil

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 437 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Conservation Service 1978). Deer, elk, and livestock Ecological Relationships and Distribution will paw through snow to find and use its semievergreen forage. Birds and rabbits use crownvetch for Richardson geranium occurs in openings through- food and cover. Crownvetch has some poisonous char- out the coniferous forest as an understory species in acteristics (Shenk and others 1976). However, when aspen and subalpine meadows (Judd 1962; USDA used in conjunction with other species, poisoning has Forest Service 1937). It is the most widely distributed geranium in North America (Hermann 1966). The not occurred (Shultz 1984). Crownvetch is normally species is not restricted to any particular soil type, but seeded as a component of a mixture to enhance forage usually occurs on fairly moist gravelly or sandy loams. production and quality. This species establishes However, it can be found on drier granitic soils and slowly, and production is low the first or second year heavy clay loams (Judd 1962; USDA Forest Service following seeding. Once established, it can withstand 1937). considerable grazing and trampling. It recovers from close grazing, but does not have the regrowth capabili- Plant Culture ties of alfalfa. When seeded in mixtures with other species, it will spread vegetatively, and will tend to Seed is generally produced yearly if plants are not dominate a community. grazed. Seed can be collected by beating it into a hopper or with a mechanical seed beater mounted on Varieties and Ecotypes a vehicle. Seed is extracted from the collected material by hammermilling or with a debearder (Stevens and ‘Emerald’, ‘Penngift’, and ‘Chemung’ have done well others 1985c). Seed can be stored for at least 5 years in mountain big sagebrush, mountain brush, and without substantial loss of viability. Acceptable ger- aspen communities. Emerald is the smallest in stat- mination is 60 percent. There are about 65,000 seeds ure and produces less forage, but it is a more aggres- per lb (143,000 per kg) at 100 percent purity. Seed can sive spreader. be cleaned to 60 percent or higher purity. Seed can be drilled or broadcast seeded on a prepared seedbed, preferably in the fall. Seeding depth should not exceed Family Geraniceae 0.25 inch (6 mm). This species can be seeded singly or Geranium richardsonii ______as a component of a mixture of other forbs and grasses. It demonstrates excellent ability to spread by natural Richardson Geranium seeding. Seeds are small and can establish on a loose seedbed or on sites where litter has accumulated. A member of the geranium family, Richardson geranium (fig. 9) is a perennial native forb, with flowers Uses and Management that are generally white with pinkish or dark veins. Stems are single or few and somewhat hairy and Forage value for cattle and sheep is rated good to viscid. Leaves are large but thin. Flowering occurs in excellent during early growth stages, and poor to good July and early August, and seed matures in Septem- later in the season. Forage value for elk (Kufeld 1973) ber. The foliage gives off a distinctive geranium odor and deer (Kufeld and others 1973), is greater than for when crushed (USDA Forest Service 1937). livestock (Hermann 1966; Judd 1962; USDA Forest Service 1937). The species develops a stout, woody root system that enables it to stabilize soil and withstand considerable grazing, trampling, and some drought. Seedings on fertile soils have been most successful. Attempting to seed or use this species on sites where it does not normally occur is not recommended. However, this forb spreads naturally and has successfully invaded roadway disturbances. Richardson geranium can persist with moderate to heavy grazing. Plants appear to be long-lived and spread well from natural seeding. Newly seeded stands should not be grazed for at least two growing seasons following planting. This species frequently increases with proper management. It persists well with seeded grasses, including smooth brome and intermediate wheatgrass. It is useful for adding diverity to mixtures, especially where shade from overstory species exists. Limited seed supplies restrict Figure 9—Richardson geranium. the use of this species.

438 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Varieties and Ecotypes There are no releases.

Family Geraniceae Geranium viscosissimum______Sticky Geranium This species occurs in the same climatic and edaphic areas as Richardson geranium. Its forage value for cattle is rated from fair to good and from good to excellent for sheep, deer, and elk (Buchanan and others 1972; Kufeld 1973; Kufeld and others 1973; Mueggler and Stewart 1980). Seed is collected, cleaned, Figure 11—Sticky geranium. stored, and seeded in the same manner as described for Richardson geranium. Seeds of sticky geranium are a little larger with about 52,000 per lb (115,000 per kg). These two species hybridize on the Wasatch Pla- teau of central Utah (fig. 10) (USDA Forest Service 1937). Sticky geranium (fig. 11) can be distinguished from Richardson geranium by its multiple, erect stems bearing deep purple to pink colored flowers, and its usually sticky and glandular herbage.

Varieties and Ecotypes There are no releases.

Family Fabaceae Hedysarum boreale ______Figure 12—Utah sweetvetch. Utah Sweetvetch Utah or northern sweetvetch (fig. 12) is a native perennial member of the legume or pea family that grows 10 to 30 inches (25 to 76 cm) tall and 30 to 80 inches (76 to 203 cm) wide. Blossoms are bright rose- pink to purple and very showy. Leaves are green above and grayish on the underside. Growth begins early in spring and provides a considerable amount of early spring forage that is highly palatable to both big game and domestic livestock. An abundance of forage is provided with some basal leaves remaining green throughout the winter (Rumbaugh 1983). This species produces a tap root (Welsh and others 1987). Some strains are rhizomatous (Ford and others 1984). This species has nitrogen fixing capabilities (Ford 1988; Ford and others 1989; Redente and Reeves 1981). Abundant seed is produced most years. Consid- erable variation occurs within the species, and there are a number of subspecies and varieties (Ford 1988; Northstrom and Welsh 1970, 1979; Rollins 1940). Utah sweetvetch is the most abundant sweetvetch in Figure 10—Hybrid of Richardson and sticky the Intermountain West. geranium.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 439 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Ecological Relationships and Distribution either machine can reduce viability of the seed lot. Seed can be stored up to 5 years without any signifi- Utah sweetvetch occurs in most Western States cant loss in viability (Stevens and Jorgensen 1994). (Ford 1988). It has fair shade tolerance (Rumbaugh Good results can be expected from direct seeding. and Townsend 1985; Wasser 1982) and can be found Seeding should occur in the fall or early winter, with in subalpine, aspen, mountain brush, ponderosa pine, the seed covered at least 0.13 inch (3 mm) but not more pinyon-juniper, and big sagebrush types. In northern than 0.38 inch (10 mm) of soil. Seeds should be inocu- Utah it has been found in the shadscale saltbush type lated with the appropriate inoculant (Redente and (Plummer and others 1968). Sweetvetch grows with Reeves 1981; Rumbaugh and Johnson 1984). Seed can 10 to 36 inches (25 to 91 cm) of annual precipitation be drilled or broadcast. Utah sweetvetch should not be on acidic or basic soils ranging from sands to heavy seeded where its seedlings are in direct competition clays (Plummer 1977), making it well adapted to with seedlings of aggressive species. If drilled, this semiarid areas of the Intermountain West (Redente forb should be seeded in rows separate from grasses. 1980). Utah sweetvetch does not produce strong seedlings. This species has a stout, branched, woody taproot, Germination is slow and seedling emergence follows thus it can take advantage of deep soil moisture, that of many other species. Seedlings and young plants resulting in considerable drought resistance and win- also develop slowly. Flowering does not occur until the third or fourth year following planting. Once estab- ter hardiness. Rhizomatous accessions have been re- lished, this species is very persistent, particularly ported (Ford and others 1984, 1989; McKell and others when seeded with other herbs. Plants can be estab- 1979; Plummer and others 1968; Rumbaugh 1984). lished by transplanting in the early spring (Institute for Land Rehabilitation 1979). Plant Culture Flowering occurs in late May and June. Seeds ma- Uses and Management ture in late July and early August. A segmented Livestock and big game make considerable use of loment or pod is formed (fig. 13) that can be harvested Utah sweetvetch (Hermann 1966; Plummer and oth- by hand or with a combine. Seed has been successfully ers 1968; Rumbaugh 1984; USDA Forest Service produced under cultivation. Seeds can be stored 1937; Wasser 1982). Spring growth provides succulent and seeded in or out of the pod. Seeds should be forage. Considerable green herbage is retained through- treated with an insecticide to control insect damage. out the growing season, and basal leaves remain green There are about 33,000 seeds per lb (73,000 per kg) at through the winter (Redente 1980; Rumbaugh 1984). 100 percent purity. Acceptable purity and germina- This species is thought to be a bloat-safe legume tion in the commercial market is 90 and 60 percent. By (Rumbaugh and Townsend 1985). It has considerable extracting the seed from the pod, unfilled, damaged, potential for improving big game and livestock ranges, and immature seed can be removed. Seed is extracted and for stabilization of disturbed and eroding areas from the pod with a debearder or Dybvig. Heat and (Plummer and others 1968; Redente and Reeves 1981). mechanical breakage resulting from improper use of Seeded areas should not be grazed for at least 2 years following seeding. Once established, this species can withstand considerable use; however, continual use has reduced its density in some areas.

Varieties and Ecotypes ‘Timp’ exhibits excellent growth rate, seed production, and nitrogen fixing capacity. This variety is fairly widely adapted.

Family Compositae Helianthella uniflora ______Oneflower Helianthella There are eight species of Helianthella in North America (Weber 1952), with oneflower helianthella having the widest distribution. Plants are upright Figure 13—Segmented loment of Utah sweetvetch. with erect heads. Seeds are flat and covered with

440 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats hair, whereas most sunflower seeds are thick and lack pubescence. Oneflower helianthella produces yellow flowers in late May and early June, and seeds mature in July (Hermann 1966).

Ecological Relationships and Distribution Oneflower helianthella (fig. 14) ranges from Montana to Oregon and south to New Mexico, mainly in mountain brush, aspen, spruce-fir, and ponderosa pine types (Hermann 1966; USDA Forest Service 1937). It does well in full sunlight and partial shade, growing on dry hillsides, open woods, and intermixed in various shrub communities.

Plant Culture Figure 14—Oneflower helianthella in full bloom. Seed matures in July and can be collected by hand, stripping, mechanical beating, or combining. Seed is Varieties and Ecotypes easily cleaned, by screening to remove debris. Seeds do not remain viable for more than 2 years (Stevens There are no releases. and Jorgensen 1994). There are just over 52,000 seeds per lb (115,000 per kg) at 100 percent purity. Family Umbelliferae Direct seeding by drilling or broadcasting has worked well. When seeding oneflower helianthella, seed should Heracleum lanatum ______be covered no more than 0.38 inch (10 mm). Germina- Cow Parsnip tion is rapid, and strong seedlings develop quickly. This species does well seeded in mixtures with other There are about 60 species of Heracleum through- herbs. Reproduction from new seedings usually out the world. However, only one, cow parsnip, occurs occurs within 3 to 4 years after the initial planting. in North America (Hitchcock and others 1961). Cow parsnip forms a compound umbel (fig. 15), with flow- Uses and Management ers occurring at the ends of long, hollow stalks. Plants are tall, often reaching 7 ft (2.1 m). The white to light The leaves, flowers, and more tender portions of the yellow flowers bloom in July and early August. Seed stems of oneflower helianthella are eaten by big game matures in late August and early September. Roots and livestock. Palatability is rated as poor to fair for are woody, short, jointed, and aromatic. Leaves are cattle and elk, and fair to good for sheep and deer divided into three very large, irregularly toothed (Kufeld and others 1973; Mueggler and Stewart 1980). leaflets, each 4 to 10 inches (10 to 25 cm) long. This perennial provides considerable forage in the spring and through the summer. Ecological Relationships and Distribution Oneflower helianthella can be seeded in mixtures with other herbs to furnish diversity and forage. The Cow parsnip is found from Alaska, eastward across plant grows on a variety of sites, occupying harsh open Canada, and south through the Western States and to conditions, yet also doing well on moist, fertile soils in Georgia. In the Intermountain West, cow parsnip some shade. When seeded with native and introduced occurs in aspen, spruce-fir, and subalpine areas, and grasses, it persists well and provides useful forage. It in meadows, with rich, loamy soils. This species prefer will spread to colonize open disturbances, and can be some shade and exists in open woodlands. used to beautify recreational sites and roadways. It usually does not provide a dense ground cover, but mixtures of oneflower helianthella with other herbs Plant Culture are useful for controlling erosion. Cow parsnip seeds are flat, obovate in shape, and New seedlings should receive at least two growing fairly large, with just over 44,000 per lb (97,000 per kg). seasons of nonuse following planting. Selective graz- Collection of seed is best accomplished by hand strip- ing of this species does not occur, and it can be easily ping or beating. If allowed to mature, good seed crops maintained in a mixture with other herbs on summer are generally produced yearly. Seeds are easily ranges. cleaned by screening to separate them from debris.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 441 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Reproduction is by seed only. Continual annual grazing will prevent seed production, reduce vigor, and result in elimination of this forb. Cow parsnip is a highly preferred species that requires deferred grazing. Seeded areas should not be grazed for at least two growing seasons following planting.

Varieties and Ecotypes There are no releases.

Family Umbelliferae Ligusticum porteri______Porter Ligusticum Porter ligusticum (fig. 16) a North American native, is a member of the umbel or carrot family (Welsh and others 1987). Plants are perennial with a characteristic aromatic taproot. White flowers are borne in a terminal umbel which is often subtended by a whorl of three to eight lateral umbels. Flower stalks are hollow and may attain a height of 4 ft (1.2 m). Flowering occurs in July and early August. Seed matures in August and September.

Ecological Relationships and Distribution Porter ligusticum is the most widespread ligusticum in the Intermountain West. It can be found in openings Figure 15—Cow parsnip in full bloom. in aspen, spruce-fir, and subalpine types and as a component of the understory in aspen stands. Porter ligusticum prefers well-drained soils. It has fairly good drought tolerance and good winter hardiness, Germination is generally low, and viability is not and can withstand considerable grazing (USDA For- maintained for much more than 2 years (Stevens and est Service 1937). Jorgensen 1994). Seed can be broadcast or drill seeded, preferably in autumn before snowfall. Seed should not be covered more than 0.5 inch (1.3 cm) deep. Seeds are rather large and are difficult to seed in mixtures with smaller seeds. Seeds must be placed in seedboxes separate from smaller seeded species so that planting rates can be properly controlled. Seeds can be easily fractured during cleaning or mechanical planting. Seedlings usually establish and grow rapidly, and can persist when seeded with most herbs. Plants normally require 3 to 4 years to attain maximum size, but once established they remain highly productive.

Uses and Management Cow parsnip is highly palatable to livestock and big game (Ellison 1951; Hermann 1966; USDA Forest Service 1937). It has considerable potential as an Figure 16—Porter ligusticum growing in a ornamental plant for summer homes, administration subalpine forb community. buildings, and recreational areas.

442 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Plant Culture Varieties and Ecotypes If allowed to mature, good seed crops are generally There are no releases. produced yearly. Seed is collected by hand stripping or beating. Seed is flat, winged, and relatively easy to separate from debris by screening. Dried seeds are Family Linaceae brittle and can be fractured or damaged by improper Linum lewisii ssp. lewisii ______cleaning or mechanical seeding. There are about 70,000 seeds per lb (154,000 per kg). Lewis Flax Seed can be stored for up to 5 years without There are approximately 100 Linum species in the appreciable loss of viability (Stevens and Jorgensen temperate regions of the world (Hitchcock and others 1994). Seed can be drilled or broadcast seeded. Fall 1961), with about 20 species occurring in the western seeding onto a disturbed seedbed has worked well if United States. Lewis flax is a delicate, erect perennial the seed is covered with soil. Seeds are heavy, irregu- sometimes reaching 3 ft (0.9 m) in height. Numerous larly shaped and do not flow well through most con- glabrous stems grow from a single crown with a woody ventional seedboxes. However, when mixed with other taproot. Flowers are generally sky blue (fig. 17), but seeds of the same size, uniform plantings can be also range from white to dark blue. The petals are achieved. Seeds are relatively large, and the gates of caducous, dropping within 1 day of flowering. Flowers most seedboxes must be opened wide enough to assure open in the morning and close by mid-afternoon passage. When this is done, seeds of smaller species (Howard and Jorgensen 1980). Flowering begins in may be planted too heavily. Consequently, this spe- May and extends through June. Spherical capsules cies is often planted separately from other seeds, and that produce up to 10 seeds each mature in late July a carrier is added to aid in uniform planting. and August. Drill seeding at rates of 2 to 4 lb per acre (2.2 to 4.5 kg per ha) in separate rows from seeded grasses Ecological Relationships and Distribution is recommended. Broadcast seeding is possible if followed by light harrowing or chaining. Although Lewis flax ranges from Alaska south to California studies have not identified specific ecotypic differ- and Texas (Harrington 1964). It is a perennial semiev- ences among populations, survival and growth rate ergreen that grows on well-drained soils ranging from differences strongly indicate distinct ecotypes exist. moderately basic to weakly acidic (Wasser 1982). Lewis Seed lots should not be planted in different elevational flax can be found in shadscale-saltbush, sagebrush zones than the collection site. Moving seed from aspen grass, and pinyon-juniper communities, and in open- communities to sagebrush types is not advised. ings in mountain brush, aspen, and conifer types, Good germination and seedling establishment usu- especially on warmer south and west exposures. This ally result from field plantings, but subsequent sur- species is intolerant of poor drainage, flooding, and vival is often low. Plants are suited to grow with other high water tables (Wasser 1982). species and persist well in mixtures once established. Some plantings have been conducted in the aspen and oakbrush communities, but this forb also exhibits usefulness in the upper sagebrush communities. It is often encountered in openings and on shallow soils, and can be seeded to improve forage conditions on somewhat harsh situations.

Uses and Management Livestock, particularly sheep, and big game make considerable use of this species in spring and summer. It is highly sought out at all dates (Hermann 1966; Kufeld and others 1973; USDA Forest Service 1937). Seeded areas should not be grazed for at least 2 years following planting. Established stands should be allowed to produce seed periodically, as reproduction is entirely from seed. Figure 17—Lewis flax in full bloom.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 443 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Plant Culture Because it maintains green basal foliage year round, this species does not burn readily, and it can be used Lewis flax can be established easily from direct as a fire suppressant species. Lewis flax has the ability seeding. Seeding can be accomplished by aerial and to compete with and spread in some cheatgrass com- hand broadcasting, drilling, and interseeding. Care munities. When seeded in mixtures, Lewis flax estab- should be taken to ensure that seeds are not covered by lishes and spreads well. It quickly occupies open areas more than 0.13 inch (3 mm) of soil. Lewis flax has been and slowly suppresses annual weeds. successfully seeded in range, wildlife, roadway, mine Flower color ranges from nearly white to dark blue. disturbance, and landscape plantings. It establishes Profuse flowering occurs for about 6 weeks (Addicott well when included in mixtures (McKenzie and others 1977), beginning in mid-May. The flowering charac- 1980; Plummer and others 1970a), and grows and teristics of this species make it a prime ornamental reproduces well in association with other species candidate. When seeded in a mixture, Lewis flax can (Stevens and others 1981b). However, young plants help make a seeding more aesthetically pleasing. can be suppressed by competitive perennial grasses. Seeded areas should not be grazed for at least 2 years This species does well when seeded in mixtures with on better sites and 3 years on drier, poorer sites. Seed shrubs. Generally, 0.25 to 0.5 lb per acre (0.3 to 0.6 kg should be allowed to mature periodically. Individual per ha) is seeded in mixtures. Reproduction is entirely plants are relatively short-lived (4 to 7 years). Abun- from seed. The species is well adapted to sagebrush, dant seed is produced when plants are not excessively pinyon-juniper, and mountain brush communities, grazed. Lewis flax will reproduce and remain in most and is one of the few forbs that can be seeded success- seedings if properly managed. fully in the salt desert types (Monsen and Plummer 1978). Everett (1982) reported that Lewis flax was the Varieties and Ecotypes superior forb in a number of pinyon-juniper seedings. It has been one of the most successful native forbs ‘Appar’ Lewis flax was released by the Utah Division seeded in the sagebrush and pinyon-juniper communi- of Wildlife Resources, the USDA Forest Service Inter- ties in Utah. Seeding is best accomplished in fall or mountain Forest and Range Experiment Station, and winter (Everett 1982; Howard and Jorgensen 1980). the USDA Soil Conservation Service Aberdeen Plant Seeds require an afterripening period to obtain maxi- Materials Center in 1980. This cultivar was named for mum germination (Eddleman 1977; Stevens and A. Perry Plummer. It was initially selected for its Jorgensen 1994). Seed can be held in storage up to 10 appearance and competitiveness with native grasses. years without any appreciable loss of viability (Stevens It is easily grown under agricultural conditions and and Jorgensen 1994). has produced up to 700 lb of seed per acre (785 kg per Seed can be collected by hand, with power-driven ha) with irrigation. ‘Appar’ exhibits a wide adaptability to sites in the Intermountain area. It is recom- beaters, or with a combine. Seed production is gen- mended for sites receiving 10 to 23 inches (25 to 58 cm) erally high. Seed is also being successfully produced of annual precipitation (Howard and Jorgensen 1980; under cultivation. Natural reproduction is usually Shaw and Monsen 1983a). Pendleton and others (1993) good, resulting in stable numbers and production report that the origin of the cultivar appears to be even though individual plants are relatively short- European and that it is derived from a naturalized lived (4 to 7 years). Flowering occurs over a 6 week population of Linum perenne. period, causing seed to mature unevenly over an ex- Maple Grove Lewis flax was released by the USDA tended period of time. There are about 280,000 seeds Forest Service, Rocky Mountain Research Station, per lb (617,000 per kg) at 100 percent purity. and the USDA NRCS Aberdeen Plant Materials Center. It is a selected germplasm of Linum lewisii Uses and Management that originated near Maple Grove, UT. Lewis flax produces basal foliage from a woody taproot (Plummer and others 1968). This species can be Family Umbelliferae classified as a semievergreen because some basal Lomatium triternatum ______foliage remains green throughout the winter. Animals paw through the snow to reach the green growth in Nineleaf Lomatium winter. Early spring growth is grazed by big game, upland game birds, and livestock. Forage value for Nineleaf lomatium (fig. 18) is a member of the livestock and big game is moderate to high, and birds umbel or carrot family (Welsh and others 1987). It and rodents seek out the seed (Everett and others was formerly identified as Lomatium simplex. It has 1978; Howard and Jorgensen 1980; Plummer and now been divided into two subspecies, L. triternatum others 1970a). Lewis flax responds well to late sum- spp. platycarpum and L. triternatum spp. triternatum mer and fall storms by producing new basal leaves. (Hitchcock and others 1961; Welsh and others 1987).

444 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Uses and Management Nineleaf lomatium is one of the first species to begin growth following snowmelt. Extremely early growth and development takes advantage of spring soil moisture and provides much needed spring succulence for big game and livestock. Deer, elk, antelope, cattle, and sheep make use of the foliage (Hermann 1966; Mueggler and Stewart 1980; USDA Forest Ser- vice 1937). Large, deep taproots enable nineleaf lomatium to withstand considerable drought, grazing, and trampling. Newly seeded areas should be given at least two seasons of growth before they are grazed.

Figure 18—Lomatium in full bloom. Varieties and Ecotypes There are no releases.

Family Umbelliferae This aromatic perennial forb, with yellow flowers and Lomatium kingii______green leaves, grows to about 14 inches (36 cm) in height. It blooms in April and May, and seeds mature Nuttall Lomatium in July and August. Nuttall lomatium, (fig. 19) formerly referred to as Ecological Relationships and Distribution Lomatium nuttallii (Welsh and others 1987), is a native perennial forb that can be found growing from Nineleaf lomatium occurs in central and southwest- sagebrush-grass up through the subalpine communi- ern Colorado, Montana, southwestern Canada, and ties. It exists on basic and acidic soils (Plummer 1977; south from California, to Colorado (Hermann 1966; Welsh and others 1987). Nuttall lomatium is not as Hitchcock and others 1961; Welsh and others 1987). widely distributed as nineleaf lomatium. It does, Nineleaf lomatium prefers well-drained or dry however, occur on drier sites and may have more rocky soils on flats, sunny open ridges, and slopes. drought tolerance. Seed is best collected by hand. L. triternatum ssp. triternatum occurs in mountain Viability is generally high. Seeds are large with ap- brush, ponderosa pine and aspen forests, and dry proximately 12,500 seeds per lb (28,000 per kg). More valleys from 5,200 to 8,500 ft (1,600 to 2,600 m). L. triternatum ssp. platycarpum is present in sagebrush- grass types, pinyon-juniper, mountain brush, ponderosa pine, lodgepole pine, and dry meadow communities from 4,300 to 9,500 ft (1,300 to 2,900 m) (Welsh and others 1987).

Plant Culture Seed matures in July and August. It can be hand collected. Screening to remove debris is all that is generally required to clean field-harvested seed. Viability of freshly harvested seed is generally good and remains high for 3 to 4 years. There are about 42,000 seeds per lb (93,000 per kg) at 100 percent purity. Seed can be drilled or broadcast seeded. Seed- ing is best accomplished in the fall, with seed being covered not more than 0.25 inch (6.4 mm) deep. This species does well when seeded in mixtures with other herbs. Figure 19—Nuttall lomatium setting seed.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 445 Chapter 19 Forbs for Seeding Range and Wildlife Habitats than 70 percent germination can be expected even after 5 years of storage (Stevens and Jorgensen 1994). Seed is collected and cleaned in the same manner as described for nineleaf lomatium. This species is readily grazed by big game during spring months. It is one of the first forbs to green up, thus providing much sought after spring and early summer succulence.

Family Fabaceae Lupinus sericeus______Silky Lupine Silky lupine (fig. 20) is a highly variable native perennial legume. Plants are usually 12 to 47 inches (30 to 120 cm) tall, growing from a branching caudex. Figure 20—Silky lupine. Flowers and foliage are usually quite abundant. Flowers are large, blue, blue-purple, or white (Welsh and others 1987). Flowers emit a distinctive pungent odor. The leaves are large, palmately compound, often Seeds are rather large with only 13,000 per lb (29,000 pubescent, and gray-green (Welsh and others 1987). per kg). Most seeds imbibe water when planted but some fail to germinate. Some seeds that fail to germi- Individual seedpods usually contain 3 to 5 seeds. nate are lighter in color than healthy seeds. Silky lupine should be seeded in the fall to meet stratifica- Ecological Relationships and Distribution tion requirements. Seed can be drilled or broadcast, Silky lupine ranges from southern Canada south to with seed covered not more than 0.25 inch (6.4 mm) New Mexico (Hermann 1966). It is common in the deep. Great Basin and can be found in sagebrush-grass, Silky lupine establishes well when seeded on disturbed or depleted sites in acid or basic soils (Plummer pinyon-juniper, mountain brush, ponderosa pine, and 1977). Fairly strong seedlings are produced; however, aspen types. Welsh and others (1987) reported that full plant development can be somewhat slow, with silky lupine is one of the most widely distributed flowers not being produced for 3 to 5 years following species in Utah, with four different varieties reported. seeding. It is most abundant in upper mountain brush and Good numbers of seedlings appear from most aspen communities. This species grows on basic, plantings. However, plant numbers are generally re- neutral, and slightly acid soils. It prefers open sun- duced quite dramatically the first growing season. light, but possesses some shade tolerance. Plants that persist the first year are usually very There are a number of perennial lupines in the hardy and will persist even under adverse conditions. Intermountain West. Most tend to intergrade with Young plants are quite competitive and persist with each other. Two common and abundant species are some competition. To date, most range seedings have silky lupine and silvery lupine. been conducted using seed collected from aspen communities. These sources are not suited to lower, drier Plant Culture elevations. Seed is usually ready to harvest in early September. Uses and Management Seed pods must be collected before they dry and seeds are shattered. Consequently, seed is collected by hand Silky lupine is one native legume that has good stripping of the complete inflorescence and, if in pure potential for revegetation of ranges in upper sage- stands, by mechanical seed beaters or combines. Fol- brush-grass, mountain brush, ponderosa pine, and lowing collection, the pods need to be further dried aspen types. This species provides early spring suc- before seeds can be extracted. During drying, collected culence and is grazed by cattle, sheep, deer, rodents, material must be covered with a screen to prevent seed and small mammals throughout the year (Hermann loss. As pods dry, they open, twist, and propel the 1966; Holechek and others 1982; Kufeld and others 1973; seed a distance of 3 to 10 ft (0.9 to 3.0 m). Lupine Plummer and others 1968). It can be seeded in mix- seeds have considerable longevity. Silky lupine seeds tures or in separate drill rows from grasses and other have been stored for 20 years and still retained 72 broadleaf herbs. It is extremely productive and pro- percent germination (Stevens and Jorgensen 1994). vides not only useful forage but also soil protection

446 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats and erosion control. It persists with grazing, and does well as an understory species. It can be used in riparian disturbances and on dry meadows that are not flooded. Silky lupine often recovers well when disturbed sites are protected for a number of years. Rodents gather and forage on lupine seeds and can interfere with seedings and natural spread. Seeded areas should not be grazed for at least 2 years, and in some cases, 3 years following planting. Once established, this species has excellent grazing and drought tolerance.

Varieties and Ecotypes There are no releases. Figure 22—‘Nomad’ alfalfa. Yellow- and blue-flowered Family Fabaceae alfalfa growing together on a pinyon-juniper site. Lupinus argenteus ______Silvery Lupine Poisonous Characteristics A major difference between silvery lupine and silky lupine is the presence of less pubescence on the leaves Some lupines, including silky and silvery, have and stems of silvery lupine. Its flowers are predomi- poisonous characteristics and can cause death if used nantly blue-purple, blue, and white. Meeuwig (1960) improperly during some seasons of the year (Davis reported that silvery lupine has outstanding forage, 1982b; Davis and Stout 1986; James and others 1980; ground cover, and revegetation potential at higher USDA Soil Conservation Service 1968). Care should elevations. Most of the perennial lupines readily hy- be taken with livestock where substantial quantities bridize (Hermann 1966). Lupinus alpestris is now of lupine are available. Livestock that obtain forage of included as part of the silvery lupine complex (Welsh other species in addition to lupine are generally not and others 1987). Seed production, seed handling, adversely affected (USDA Soil Conservation Service seeding, and seedling establishment characteristics of 1972b). When planting lupine, care should be taken to silvery lupine are similar to those of silky lupine. ensure that it is seeded in mixtures with other spe- Silvery lupine occurs at higher elevations than does cies, and that large, solid stands are not established. silky lupine. Major areas of occurrence are aspen, spruce-fir, and subalpine herblands (fig. 21). Family Fabaceae Medicago sativa Medicago falcata ______Alfalfa Sicklepod Alfalfa Two species, Medicago sativa (alfalfa) and M. falcata (sicklepod alfalfa), were introduced to North America (Hansen 1913). Alfalfa is generally blue-flowered (fig. 22) and nonrhizomatous, whereas sicklepod alfalfa is generally yellow-flowered and somewhat rhizomatous. Both species are perennials. Stems normally reach 11 to 40 inches (30 to 100 cm), depending on growing conditions. Stems are branched; some are ascending. Most range varieties form decumbent stems at the base of the plant. Varieties cultivated for hay produc- Figure 21—Silvery lupine in a subalpine community. tion or irrigated pastures are normally more erect.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 447 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Leaves of alfalfa are glabrous to hairy, alternate, and pinnately trifoliolate; the leaflets are serrate distally and oblanceolate to oblong. Flowers are small, usually less than 7 to 10 mm long. Fruits consist of several-seeded spiral pods (Harrington 1964). Plants produce an abundance of herbage, with growth beginning early in the spring. Alfalfa is a complex taxon with nine recognized subspecies (Gunn and others 1978). These express intergrading morphologies and hybridize quite freely. The genus evolved in the Mediterranean region. The perennial mesophytic subspecies arose in Western and Central Asia (Lesins and Lesins 1979). Diploid (2N = 16) and tetraploid (2N = 32) forms commonly occur. Most cultivars are tetraploid (Rumbaugh and Townsend 1985). Lesins and Lesins (1979) report that Figure 23—‘Rambler’ alfalfa seeded into rootplow the diploid populations of alfalfa originate from steep strips in a Gambel oak-mountain big sagebrush mountainous regions. community. Miller and Melton (1983) described over 400 cultivars and strains of alfalfa in North America. Today there are many more. The origin of the 15 most commonly grazed populations are described by Seedling establishment is rapid, and seeding suc- Rumbaugh (1982a). All dryland forage types are inter- cess rate can be high. Grazing stimulates rhizome specific hybrids of alfalfa and sicklepod alfalfa production, which results in increased crown size and (Rumbaugh and Townsend 1985) (fig. 23). These ground cover (Rosenstock and Stevens 1989). authors reported that sicklepod alfalfa contributes Alfalfa is well suited to harsh sites and infertile genes that increase drought tolerance, winter hardi- soils. Under these conditions seedlings of few other ness, and tolerance to grazing. Alfalfa germplasm herbs survive as well as alfalfa. Once established, enhances resistance to disease and insects and im- alfalfa serves as an excellent nurse crop. Although proves forage and seed production. plants begin growth early in the spring, they do not Most range and field varieties of alfalfa used in prevent the entry of other species. Alfalfa may produce Western range plantings arose from the early efforts considerable annual litter that provides a good cover of Dr. N. E. Hansen (1913) of South Dakota. Early for the seedbed. Once established, alfalfa is very per- testing conducted by Dr. Hansen, led to identification sistent (Rumbaugh 1982b) (fig. 25), but little repro- of population with spreading root systems. This char- duction has occurred from established stands on acteristic has subsequently been bred into field and rangeland sites (Rosenstock and Stevens 1989). Rhi- range varieties (Rumbaugh and Pedersen 1979). zomatous forms do spread vegetatively even under Canadian scientists later advanced the usefulness of arid conditions. this species by breeding varieties with better grazing and cold tolerance (Heinricks 1963). Alfalfas were entered into extensive field plantings within the Intermountain region in the 1930’s. Numerous field and range varieties have subsequently been developed.

Ecological Relationships and Distribution Rangeland varieties of alfalfa are well adapted to areas that receive 10 inches (25 cm) or more of annual precipitation. They are especially well-adapted to sagebrush/grass, pinyon-juniper, and mountain brush (fig. 24) communities. They have been seeded and have performed fairly well in aspen, spruce-fir, and subalpine types. This forb is semievergreen, having green basal leaves throughout the winter. Growth starts in the spring when the plant is exposed from underneath the snow. Once grazed, regrowth of rangeland types is limited compared to that of irrigated Figure 24—‘Ladak’ alfalfa on a 27-year-old alfalfas. pinyon-juniper chained site.

448 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

seedboxes and eliminate seed separation when mixtures are planted. Although alfalfa seeds should be planted slightly shallower than large seeded grasses, adequate stands normally appear when they are drilled together. Al- falfa seedings develop rapidly and compete favorably with many grasses and other broadleaf herbs. Alfalfa seedlings, however, do not compete well with seedlings of crested and intermediate wheatgrass and smooth brome. Because of the possibility of bloat, alfalfa should be seeded as a component of a mixture when planted in a pasture (Lorenz 1982). Up to 40 percent by weight of a grass-legume mixture has been seeded to alfalfa without causing bloat problems to livestock. Bloat has Figure 25—Rehabilitated range 21 years fol- not been observed as a problem on mixed rangeland lowing chaining of juniper-pinyon and aerial seedings. seeding. Thirteen grasses, shrubs, and forbs Because alfalfa is highly utilized by livestock, big were seeded. After 21 years alfalfa is the and small game, birds, rodents, and rabbits, a suffi- predominant forb. cient number of plants must be established to ensure the species is not eliminated by overuse. On most ranges, at least 2 lb of seed per acre (2.2 kg per ha) should be planted in the seed mix. Seed costs are Plant Culture generally not prohibitive, and under some conditions alfalfa should be planted at even higher rates. As Alfalfa is the most widely seeded rangeland forb in seeding rates are increased, stand density also in- the Intermountain West (Rumbaugh 1984; Rumbaugh creases. Seeding alfalfa at rates exceeding 6 to 8 lb per and Townsend 1985). Depending on the rangeland acre (6.7 to 9.0 kg per ha) is unnecessary. There are sites and seeding technique, it can be seeded in the fall about 220,000 seeds per lb (485,000 per kg). Purity is or spring. Planting early in the spring usually pro- generally over 95 percent with germination at least duces good stands, but plantings in late fall or winter 85 percent. Under warehouse storage seed longevity is have been satisfactory in large restoration programs. excellent and little viability is lost after 25 years Late winter plantings may prevent precocious ger- (Stevens and Jorgensen 1994). mination (Plummer and others 1968), but alfalfa Alfalfa is able to fix nitrogen, thereby improving germinates quickly in early spring and may succumb soil fertility. Johnson and Rumbaugh (1981) re- to spring frosts. ported that alfalfa appears able to fix nitrogen even Seed can be drilled or broadcast seeded, but it must during periods of drought when other legumes do not be covered. Aerial seeding followed by anchor chaining nodulate. or pipe harrowing and aerial seeding on snow over disturbed soil has produced excellent rangeland stands. Uses and Management Broadcast planting has often been more successful Alfalfa is heavily grazed by livestock, big and small than drilling (Plummer and others 1968). Fall aerial game, upland game birds, rodents, and rabbits during seeding on a rough seedbed has resulted in excellent all seasons (fig. 26). The inclusion of alfalfa in a stands (fig. 25). Whether broadcast or drilled, seed rangeland seeding can have a number of positive should be covered 0.25 to 0.5 inch (6 to 13 mm) deep effects. It can (1) increase total herbage production, (2) (Plummer and others 1968; USDA Soil Conservation increase production and protein content of associated Service 1971a). species (Johnson and others 1983; Rumbaugh and Alfalfa is particularly adapted to drill or broadcast others 1981, 1982), (3) extend the grazing season, both seeding in mixtures with other species. Seeds are during spring and fall, (4) increase diversity (flora and rather small, round, and smooth. When mixed with fauna) of seeded communities, and (5) improve soil grasses, seed sorting and separation can occur in the stability. Species growing in association with alfalfa seedbox. Alfalfa seed usually does not separate when are generally positively affected by its ability to fix mixed with grasses if planted at a ratio of 1 to 5. nitrogen (Johnson and others 1983; Rumbaugh and Separate legume seedboxes are mounted on most others 1981, 1982). Consequently, alfalfa can be drills. The legume boxes are designed to dispense the interseeded into established grass stands to improve small alfalfa seed more accurately than the larger vigor and herbage production.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 449 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Peterson (1979) reported excellent stands persist 23 to 28 years after seeding on sites with 8 to 11.5 inches (20 to 29 cm) of precipitation. The Utah Division of Wildlife Resources has aerial seeded alfalfa on over 150,000 acres (60,700 ha) of game range during the past 30 years. At a few sites, there has been a major loss in plant density. Where a decrease in density has occurred, it has been a result of continuous heavy rabbit use and continual spring use by cattle. Similar results have been reported by Rumbaugh and Pedersen (1979) and USDA Soil Conservation Service (1971a). Rumbaugh (1982b) reported that stand density of eight different varieties was adequately maintained during a 25-year period, with proper spring grazing by cattle. Where insufficient seed is planted and plant Figure 26—Alfalfa seeding in a chained pin- density is low, considerable grazing pressure will yon-juniper basin big sagebrush area. Deer reduce the stand At least 2 lb. (0.9 kg) of seed per acre use only on left, rabbit and deer use on right. should be seeded to ensure sufficient density to sustain grazing. Alfalfa generally receives more grazing pressure than other associated species and withstands greater use Alfafa is compatible with native species in seedings than other broadleaf herbs. Weakened stands recover in mountain brush, pinyon-juniper and big sage- readily when grazing pressures are removed. The deep brush communities. The forb has not noticeably di- taproot, large crowns, and extensive rhizomes enable minished or eliminated native broadleaf herbs or the plant to persist over extended periods of drought. grasses. New seedings can be damaged by continuous grazing Regrowth capacity varies among varieties. Deep- by rabbits, rodents, livestock, or big game animals. rooted types possess better regrowth capabilities than Insects, including grasshoppers, have seriously defoli- rhizomatous range types. Alfalfa initiates growth early ated alfalfa, often within a 2 to 5 day period. Attempt- in the spring and remains green and productive as ing to seed alfalfa and palatable grasses in areas with long as soil moisture permits. It can recover following high rabbit populations can be risky. Once estab- grazing or clipping. It normally attains maximum lished, mature plants can usually recover from cyclic production after most cool-season grasses cease periods of heavy use, but young seedlings cannot. growth. Certain strains of grass, including ‘Latar’ Alfalfa can be used to sustain big game animals orchardgrass and ‘Greenar’ intermediate wheatgrass, during the early spring and summer months. Plant- mature when alfalfa is at the optimum stage of growth ings established on pinyon-juniper sites in central for harvesting as hay (Hafenrichter and others 1968). Utah clearly attract deer and provide a substantial These species can be planted in mixtures to signifi- part of their diet, beginning in early March and con- cantly extend the season of grazing and enhance total tinuing until early June (Rosenstock and others 1989). herbage production. Alfalfa also furnishes excellent Wildland sites seeded to support big game should be fall green-up, and standing crops cure quite well. grazed only lightly by livestock. If plants are subjected Consequently, this species has been used in wildlife to stress by drought or excessive use by insects or seedings to attract and keep game animals from tres- rabbits, livestock grazing should be curtailed. passing in agricultural fields. Areas seeded to alfalfa Alfalfa has been seeded on some rather arid sites, should not be grazed for a minimum of two growing including sites dominated by Wyoming big sagebrush seasons following seeding. Early use can result in poor and shadscale saltbush. When seeded during years of survival and loss of vigor (Berdahl and others 1986). average precipitation, plants usually establish. How- Drier areas may require additional time before stands ever, failures can be expected if seeding occurs during fully establish. years with low precipitation. Plantings can be main- It is not unusual for seedlings and young plants to tained under arid circumstances if grazed judiciously. succumb during the first 1 to 3 years following planting (Rosenstock and Stevens 1989). Once established, Varieties and Ecotypes however, alfalfa has considerable longevity, resistance to grazing and trampling, and moderate shade A number of strains of alfalfa—‘Ladak’, ‘Rambler’, tolerance. Kilcher and Heinrichs (1966b), Pearse (1965), and ‘Nomad’—are adapted to ranges where annual Rosenstock and Stevens (1989), and Rumbaugh and precipitation exceeds 10 inches (25 cm) (Plummer and

450 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats others 1968; Rumbaugh 1982b). These strains produce much larger crowns than the usual field strains and have more complex root systems. In addition, they survive better from underground attacks by gophers and other rodents. Some plants also spread by stem layering. Consequently, these strains are superior for seeding rangelands (Plummer and others 1968). ‘Ladak’ is the best known and most widely used rangeland variety. If seeds of several strains are available, a mixture may be used. The varieties mentioned above have shown unusual persistence and production under heavy grazing and lower precipitation (Plummer and others 1968). Other cultivars—‘Spreader’, ‘Spreader II’, ‘Rhizoma’, ‘Runner’, ‘Travois’, ‘Teton’, and ‘Drylander’—have also demonstrated good adaptability to wildland sites. Additional varieties are currently being developed for rangelands. A major concern is to improve drought tolerance so the forb may be planted in more arid environments.

Family Fabaceae Melilotus officinalis ______

Yellow Sweetclover Figure 27—Yellow sweetclover. The two most abundant sweetclovers on Western ranges are yellow sweetclover (yellow-flowered), and white sweetclover (white-flowered). Neither are native to North America. There are annual and biennial Plant Culture forms of each; however, most populations are bienni- als (Rumbaugh 1984; Smith and Gorz 1965). During Seed production of yellow sweetclover is gener- the first season, only vegetative growth occurs; during ally high. There are more than 250,000 seeds per lb the second season, long, well-branched stems 1 to 6 ft (550,000 per kg). Germination is generally high, and (0.3 to 1.8 m) tall (fig. 27) develop. Leaves are trifoli- good viability is retained for at least 2 years with olate, and dentate margined, with small narrow warehouse storage. Adequate, inexpensive seed is stipules. Flowering occurs on long narrow racemes, generally available. Like most legumes, sweetclover and single seeds are produced in straight smooth seeds have a hard coat and are best seeded in the fall pods (Hermann 1966). After flowering and seed or winter. Seeds can lie dormant in the soil for years maturation, plants usually die. and then germinate when temperatures and moisture conditions are favorable. In central Utah, seedings Ecological Relationships and Distribution more than 35 years old still retain some yellow sweetclover. Nichols and Johnson (1969) report that Yellow sweetclover originated in Europe and Asia with 15 inches (38 cm) of precipitation, yellow and is now distributed over most of the United States sweetclover will reseed successfully. Seeds can be and Southern Canada. The species is especially adapted drilled or broadcast on relatively unprepared seed- to disturbed sites. It has a strong seedling, fairly good beds. Best stand establishment occurs when seeds are drought resistance, and good winter hardiness (USDA planted 0.13 to 0.25 inch (3 to 6 mm) deep, preferably Forest Service 1937; USDA Soil Conservation Service on a fairly firm seedbed. 1971b). It grows well with over 10 inches (25 cm) annual precipitation in aspen, mountain brush, Uses and Management ponderosa pine, big and black sagebrush, and pinyon-juniper types. Yellow sweetclover also does Yellow sweetclover is frequently seeded on Western well in saline and alkaline soils and on many riparian rangelands (Rumbaugh and Townsend 1985). It is an sites (Plummer 1977; USDA Soil Conservation Ser- effective erosion control species because it has an vice 1968, 1971b; Wasser 1982). extensive, deep taproot system and a strong vigorous

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 451 Chapter 19 Forbs for Seeding Range and Wildlife Habitats seedling. It also produces a dense ground cover in 1 or 2 months and fixes nitrogen. Up to maturity, palatability is high for big and small game (Graham 1942; Kufeld 1973; Kufeld and others 1973; Mueggler and Stewart 1980; Plummer and others 1968; Rumbaugh 1984; USDA 1971b) and for livestock (Hermann 1966; USDA Forest Service 1937; USDA Soil Conservation Service 1971b), with forage quality reportedly higher than that of alfalfa (Magness and others 1971; Rumbaugh 1984). Waterfowl, upland game birds, and bees also use yellow sweetclover (Autenrieth and others 1982; Rumbaugh 1984; USDA Soil Conservation Service 1971b). The species has a very strong seedling that emerges in the spring, providing early succulence. Yellow sweetclover provides much needed live ground cover and forage during the critical first and Figure 28—Tall bluebell in full bloom. second years. It can be seeded at 0.5 to 2 lb per acre (0.56 to 2.2 kg per ha), (Nichols and Johnson 1969; Plummer and others 1968; USDA Soil Conservation 11,000 ft (1,800 to 3,400 m). The species can be found Service 1971b). Yellow sweetclover is generally seeded along streams in canyon bottoms, meadows, parks, with perennials on disturbed sites. It furnishes initial and in scattered timber stands. It grows well in shade cover and forage as slower developing herbs attain and open situations in pure stands as well as in mixed mature stature. It withstands heavy use and can communities. Tall bluebell is present in all mountain reseed following fires or other disturbances. regions of Utah, southwestern Wyoming, and northwestern Colorado (Cronquist and others 1984). Varieties and Ecotypes Plant Culture There are three released varieties—‘Goldtop’, ‘Madrid’, and ‘Yukon’ (Wasser 1982). These varieties Flowering occurs during midsummer. Seeds mature have not performed any better than the common seed in September and can be collected by hand stripping, sources. Most seed marketed is from uncertified beating, and in pure stands with various types of common lots. combines. Germination is generally high (60 to 70 percent), with good viability being maintained for at Family Boraginaceae least 5 years. Seeding is best accomplished in the fall on disturbed soil, with seed covered 0.25 to 0.5 inch Mertensia arizonica ______(6 to 13 mm) deep. Tall bluebell can be seeded in mixtures with other herbs. It usually grows on fertile Tall Bluebell soils. When planted on disturbed, degraded sites it has Tall bluebell is the most widely distributed Mertensia not done well. Under good conditions, plants establish in the Intermountain West (Higgins 1972; Mathews quickly and reach maturity in 3 to 5 years. Plants 1968). There are, however, other bluebells that are spread slowly by rhizomes, yet are very persistent and locally important (Hermann 1966). Different varieties compatible with other native species when seeded in of tall bluebell have been described, particularly from mixtures. collections obtained in Utah (Cronquist and others 1984; Welsh and others 1987). Tall bluebell is a rhi- Uses and Management zomatous native perennial with a thick, woody root. It Sheep, elk (Kufeld 1973), and deer (Kufeld and grows to a height of 2 ft (61 cm) and produces an others 1973) seek tall bluebell and gain weight excep- abundance of pale blue to very dark blue flowers tionally well on the herbage. Cattle prefer the flower (fig. 28). Leaves are abundant and are borne on erect stalks but will make use of all plant parts (USDA stems. Fruits are hard and consist of four nutlets Forest Service 1937). Sheep graze the herb whenever (Cronquist and others 1984). available. The bluebells are generally very showy and fairly easily seeded or transplanted, and are good soil Ecological Relationships and Distribution stabilizers. Consequently, the plant is a good candi- Tall bluebell occurs in fairly moist to well-drained date for use in recreational areas, summer home sites, soils in aspen, spruce-fir, ponderosa pine, and sub- and areas with high aesthetic value and use. The alpine types, usually at elevations from 6,000 to plant furnishes such excellent forage that it should

452 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats be included in most seedings on adapted sites. How- inches (36 cm) of annual precipitation. Sainfoin pre- ever, the lack of sufficient seed currently curtails the fers well-drained, deep calcareous soils (Eslick 1968), use of this forb. This species has not done well if exhibits fairly good salt tolerance (Jensen and Sharp planted off site, particularly at lower elevations where 1968), and grows well in soils low in phosphorus it normally does not occur. (Rumbaugh and Townsend 1985). Newly seeded areas should not be grazed for at least 2 years. Large fibrous root systems, once established, Plant Culture allow tall bluebell to withstand periods of drought, Single seeds are produced in pods and are marketed grazing, and trampling. Concentrated use can occur if unshelled. There are about 26,000 unshelled seeds seedings are not properly managed. per lb (57,000 per kg). Unshelled seeds are best drilled, Varieties and Ecotypes but can be broadcast if covered sufficiently. Seed should be covered at least 0.25 inch (6 mm) and not There are no releases. more than 0.75 inch (19 mm) deep (Jensen and Sharp 1968). Seeding is best done on a firm seedbed that is relatively weed free. For range seeding, it is best to Family Fabaceae seed in the fall. Seed should be inoculated with the Onobrychis viciaefolia proper rhizobium (Dubbs 1968). Seed is marketed with about 70 percent germination and 95 percent Sainfoin purity. When seeded in a mixture, about 2 to 5 lb per acre (2.2 to 5.6 kg per ha) is recommended. Seed can Sainfoin (fig. 29) is a perennial, nonbloating, intro- be stored up to 4 years without much loss of viability. duced legume that grows 1 to 3 ft (30 to 91 cm) tall, and This species can be seeded individually or as a produces white or pink to purplish pea-like flowers. Stems are semierect, originating from a branched root component of a mixture. A strong seedling is usually crown attached to a deep main taproot. Leaves are produced. Rate of growth following seeding is only fair alternate, odd-pinnate and oblong to elliptic or oblan- on mesic sites. However, plants are quite persistent ceolate (Rumbaugh and Townsend 1985; Welsh and and hardy. Two years are required for flowering to others 1987). occur under favorable conditions.

Ecological Relationships and Distribution Uses and Management Sainfoin was introduced into North America from Sainfoin is nonbloating and palatable to livestock, Europe. It has been used primarily as a hay or pasture big game, and sage-grouse (Autenrieth and others crop. However, the species demonstrates considerable 1982; Holder 1968; Rumbaugh 1984; Shaw and Coo- usefulness for rangelands. per 1973; USDA Soil Conservation Service 1968a). Sainfoin is adapted to sagebrush, pinyon-juniper, Mule deer in Utah have been observed to prefer sain- and mountain brush areas that receive at least 14 foin hay over alfalfa hay. Sainfoin has a deep taproot and has wilt and drought hardiness equal to or slightly less than does ‘Ladak’ alfalfa (Dubbs 1968; USDA Soil Conservation Service 1968a). Some basal leaves remain green throughout most of the winter. Sainfoin starts growth early in the spring, usually prior to alfalfa. It continues to grow or remains green throughout the summer period. Once grazed or clipped, regrowth is limited. Characteristics that give sainfoin considerable potential in the Intermountain area as a forage crop include early greenup, cold and drought tolerance, nonbloating characteristics, high forage production potential, ability to grow with other species, and adaptation to calcareous soil. Areas seeded to sainfoin should not be grazed for at least the first two growing seasons. Every 2 to 3 years, seed should be allowed to mature and seed naturally. Plants are persistent and withstand heavy clipping or Figure 29—Sainfoin growing with crested wheat- grazing. Once established, this species also competes grass and basin big sagebrush. fairly well in mixed communities.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 453 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Varieties and Ecotypes A number of improved varieties have been released, including ‘Eski’, ‘Melrose’, ‘Nova’, ‘Onar’, ‘Remont’, and ‘Runemex’. All were released for cultivated pastures and for hay. ‘Eski’ is the most commonly seeded variety on ranges. This variety does well when seeded in upper pinyon-juniper, mountain big sagebrush, and mountain brush areas.

Family Umbelliferae Osmorhiza occidentalis ______Sweetanise Sweetanise is an erect, perennial, aromatic native that grows up to 4 ft (1.2 m) tall (Welsh and others 1987) (fig. 30). Crushed leaves and seed give off a Figure 30—Sweetanise with mature seed. heavy licorice odor (Hermann 1966). It is a member of the umbel or carrot family and produces small umbels of yellowish-green flowers. Flowering occurs in May Uses and Management and June, and seed matures in September. Fruits are slender and sharp-pointed. Cattle, sheep, and big game show a particular fond- ness for the foliage and for developing seeds of Ecological Relationships and Distribution sweetanise (Hermann 1966; Kufeld 1973; Kufeld and others 1973; Plummer and others 1955; USDA Forest Sweetanise occurs from Western Canada south Service 1937). The plant is palatable and remains through Utah and Colorado. It grows well in shade, green throughout the grazing season. Consistent seed open areas, mixed communities, and in solid stands. crops are produced yearly if seeds are allowed to Sweetanise prefers cool, moist woods, moist hillsides, mature before being grazed. Large, thick, woody roots valleys, and forest openings in aspen, spruce-fir, and allow the plant to withstand considerable use during subalpine types. It grows on basic, neutral, and slightly dry periods. This species does not furnish a dense acidic soils. ground cover and is not recommended for erosion control. Heavy grazing can eliminate or reduce its Plant Culture density. Seeded areas should not be grazed for at least 2 This species can be successfully established by di- years following planting. A fairly strong seedling is rect seeding. Lack of a consistent seed source reduces produced, and the rate of growth is good. Seeds are seeding of this species. It establishes easily and spreads generally produced the second year following plant- quickly from seed. Seeding is best completed in the ing. Maturing seeds are readily eaten by grazing fall. Seed should be covered, but not more than 0.25 animals. Some type of deferred grazing should occur to inch (6 cm). Sweetanise does well seeded in mixtures. allow seeds to mature at least every 2 to 3 years. Seed is usually collected by hand, but dense stands can be mechanically harvested. Seeds are brittle and can Varieties and Ecotypes be damaged during cleaning and seeding. Seeds can generally be cleaned by screening from harvested There are no releases. debris. There are about 30,000 seeds per lb (73,000 per kg). Seed should not be stored more than 2 years, or Family Scrophulariaceae considerable loss in viability can occur. Thirteen year old seed is reported to exhibit 44 percent germination Penstemon palmeri ______(Stevens and Jorgensen 1994). Seeds are large and Palmer Penstemon usually must be drilled seeded to ensure proper soil coverage. Seeds are approximately the same length as In North America there are over 200 native spe- some grass seed and can be planted in a mixture, cies of Penstemon (Hermann 1966; USDA Forest Ser- depending on the species sown. vice 1937). Many are short-lived herbaceous species

454 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats that spread readily by seed. A number are rhizomatous. Some penstemon are semievergreen, producing basal leaves that persist the entire year. They can be found in most vegetative types of Western North America. Palmer penstemon (fig. 31) has long flowering stalks 20 to 55 inches (50 to 139 cm) tall with large showy pink blossoms that are produced in late spring and early summer. Seed matures in September. Flowers give off a pleasant fragrance that is unique among the penstemons, as they usually have little or no scent (Stevens and others 1985b).

Ecological Relationships and Distribution Figure 32—Broadcast seeding of Palmer pen- Palmer penstemon is an evergreen native perennial stemon on a roadside disturbance. that occurs in blackbrush, sagebrush-grass, pinyon- juniper, mountain brush, and ponderosa pine types of the Intermountain West. It is a fairly short-lived (4 to 5 years) pioneering species. Plants spread by Plant Culture seed. This species can be found growing on basic and acidic soils (Plummer 1977). It produces long seed Good results can be expected from drilled or broad- stalks and an abundance of basal evergreen leaves cast seedings on disturbed soils and on raw and that usually remain gray-green throughout the year. eroding sites (fig. 32) where it occurs naturally as an A large fibrous root system is produced. Palmer pen- early invader. Seed can be drilled or broadcast onto stemon has considerable grazing tolerance and good disturbed soil, with covering not to exceed 0.15 inch drought tolerance. (3 mm). Natural reproduction generally compensates for the short-life span of individual plants. Seed is generally produced in abundance. Some dormancy exists that allows seeds to accumulate in the soil until favorable conditions occur for germination and establishment. Seeds of most penstemons, particularly Palmer penstemon, can be seeded in mixtures with most other herbs. Small vigorous seedlings appear early in the spring. They compete well and usually are not eliminated by competition from other species. When seeded heavily, considerable thinning or dieoff of young seedlings normally results. However, sufficient survival usually occurs to provide adequate stands. Palmer penstemon is particularly adapted to harsh situations and tends to dominate or establish on areas where other species fail. It is one of the few native broadleaf herbs that most consistently establishes from seeding. It is particularly important for its ability to establish and enhance the establishment of other plants on harsh sites. All species of penstemon have small seeds that are easily cleaned to a purity that exceeds 90 percent. Palmer penstemon has 350,000 seeds per lb (770,000 per kg) at 100 percent purity. Seeds are so small that they are difficult to meter through most conventional drills. If seeded alone, a carrier is often used to regulate planting rates. Seeds can be mixed with seeds of most other species although some separation can occur in the seedbox. When seeded in mixtures, 1 to 3 lb per acre (1.1 to 3.4 kg per ha) are Figure 31—Palmer penstemon in full bloom. adequate.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 455 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Seeds can be collected by hand stripping or beating; pure stands can be combined. Seeds are cleaned by hammermilling and screening. Storage of up to 5 years has resulted in no significant loss in germination (Stevens and Jorgensen 1994). Seeds are being produced commercially.

Uses and Management Palmer penstemon is eaten readily by deer and elk, especially during winter and spring months (Kufeld and others 1973; Plummer and others 1968). Basal leaves are green and succulent all year long. Cattle and sheep also make use of this species. The large, Figure 33—Wasatch penstemon in a mixed sub- fibrous root system and large, succulent basal leaves alpine forb community. provide considerable soil protection. This species is an early invader and does especially well on disturbed raw soils. Pink blossoms in late spring and early summer however, somewhat larger with about 230,000 per lb make this plant useful for beautification projects. (510,000 per kg) at 100 percent purity. This species is Because of seed dormancy, seeding should occur in particularly important as a forage and cover plant, the fall. Once established, stands are self-perpetuat- and establishes readily from artificial seeding. It pering. Seeded areas should not be grazed for at least two sists under heavy grazing and can be seeded in mix- growing seasons following planting. Flowering and tures with other herbs. seed production often occur the second year following planting. This species has little shade tolerance, but grows as an understory with sagebrush. It is some- Family Scrophulariaceae what resistant to fire and trampling, but can be elimi- Penstemon eatonii ______nated by excessive grazing. Eaton or Firecracker Penstemon Varieties and Ecotypes Eaton or firecracker penstemon (fig. 34) is a bright ‘Cedar’ was selected and released in 1985 (Stevens red to scarlet-flowered perennial semievergreen na- and others 1985b) for its wide area of adaptation, tive that occurs primarily on rocky soils. It occurs in winter succulence, forage and seed production, and grazing preference by livestock and wildlife. ‘Cedar’ does well in blackbrush, salt desert shrub, sagebrush- grass, pinyon-juniper, and mountain brush types within the Intermountain region. Commercial seed is readily available and recommended for use.

Family Scrophulariaceae Penstemon cyananthus ______Wasatch Penstemon Wasatch penstemon (fig. 33) is a native perennial with bright semievergreen basal leaves. Plants develop from a primary taproot and produce several 2 to 3 ft (61.0 to 91.4 cm) long stems with deep blue to nearly purple flowers. Wasatch penstemon can be found on rocky soils, but is usually associated with more fertile, deeper soils that may be slightly acidic or basic. It occurs from the mountain brush type to higher elevations, including the subalpine zone. Flowering occurs from May through July. Seeds are formed in September. Seed processing and planting features are similar to those of Palmer Figure 34—Eaton or firecracker penstemon and most other penstemons. Seeds are, penstemon.

456 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats mixed desert shrubs, sagebrush-grass, mountain brush, and aspen communities. This species forms a taproot. It is generally found on harsh, rocky sites; but it also occurs on more favorable sites. Seed collection, handling, and seeding techniques are similar to those described for Palmer penstemon. Seeds can be stored up to 12 years without much loss of viability (Stevens and Jorgensen 1994). Eaton penstemon is particularly useful for seeding in pinyon-juniper, mountain brush, and big sagebrush sites. It is not an abundant seed producer but establishes easily and is a useful forage species. Its brilliant flowers are extremely attractive and useful in landscape plantings. ‘Richfield’, a source-identified variety, has been released.

Family Scrophulariaceae Figure 35—Rocky Mountain penstemon. Penstemon humilis ______Low Penstemon

This semievergreen native penstemon is a low- soils. Rydberg penstemon has a dark violet-purple growing, 12 inch (30 cm) tall perennial forb with a flower and dark green basal leaves that are present matted, much-branched root system. Flowers are throughout the year. It is highly sought by sheep and borne on short flower stalks and range in color from big game animals. Seed and seeding characteristics deep-blue to violet-blue. Foliage is light green. Low are similar to those of Palmer penstemon. penstemon can be found in sagebrush-grass, pinyon- juniper, mountain brush, aspen, and subalpine communities, generally on gravelly, well-drained soils. Family Scrophulariaceae Planting procedures and seed collection are similar to those of other penstemons. This penstemon is eaten Penstemon strictus ______by wildlife and livestock and can be used to stabilize Rocky Mountain Penstemon erosive sites. This is a semievergreen penstemon that has an Family Scrophulariaceae abundance of dark, shiny green leaves and blue to violet flowers borne on stalks 1 to 2.5 ft (30 to 76 cm) Penstemon pachyphyllus ______tall. Rocky Mountain penstemon (fig. 35) does well on Thickleaf Penstemon rocky and sandy loam soils that range from weakly acidic to alkaline. This species does best with 15 to Thickleaf penstemon, as its name indicates, has 20 inches (38 to 50 cm) of annual precipitation. Seed- broad, fleshy, thickened, bluish-green leaves covered ing and seed handling requirements are similar to with wax. Flowers are light purple to bluish-violet. those of Palmer penstemon. One variety, ‘Bandera’, Seeds are small with 335,000 per lb (739,000 per kg) at has been released, and seed is being produced com- 100 percent purity. Seeds can be stored for up to 14 mercially. ‘Bandera’ was selected for its longevity and years without appreciable loss of viability (Stevens excellent forage and seed production. It is well adapted and Jorgensen 1994). Seeding requirements are simi- to mountain brush, ponderosa pine, and spruce-fir lar to those of Palmer penstemon. This species occurs areas. in eastern and northern Utah in salt desert shrub, sagebrush-grass, pinyon-juniper, mountain brush, and conifer communities. It is eaten by wildlife and live- Family Rosaceae stock, and provides excellent soil protection. Sanguisorba minor ______Family Scrophulariaceae Small Burnet Penstemon rydbergii ______A member of the rose family, small burnet (fig. 36), Rydberg Penstemon is an introduced perennial with a basal rosette of pinnately compound leaves arising from a caudex and This species occurs in mountain brush, aspen, conif- taproot. Numerous flowering stalks may grow from erous forests, and open parklands on basic and acid 2 to 20 inches (5 to 51 cm) in height. Flowers are

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 457 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

covered, but not more than 0.25 inch (6 cm) deep. Fall and winter seedings have proven to be the most successful. Because of the rapid rate of germination and seedling development, small burnet can usually be successfully established by spring seedings. Small burnet has been seeded widely throughout the West (Plummer 1977; Plummer and others 1970a,b; Stevens and others 1977). Following germination, growth is rapid, resulting in good ground cover within 1 or 2 years. A large amount of foliage and seed is usually produced, even during the first year of growth. Small burnet is a excellent species to seed in mixtures. Seed lots of small burnet germinate between 80 and 95 percent, and retains high viability when stored for over 25 years (Stevens and Jorgensen 1994). Seeds Figure 36—Small burnet. undergo afterripening, with germination improving yearly up to 3 years following harvest (Stevens and Jorgensen 1994). There are 55,000 seeds per lb (120,000 per kg) at 100 percent purity. Seed is marketed at formed in dense, terminal heads. Basal leaves remain about 95 percent purity and 90 percent germination. green nearly the entire year. Growth starts in early Seedling emergence and establishment for small bur- spring, flowers appear in May and June, and seed net is high (5 to 8 percent) compared to many other matures in August and early September. range species (Everett 1982). Most plants appear to have a lifespan of 7 to 12 years, Ecological Relationships and Distribution but some have persisted to 20 years (Plummer and others 1968; Stevens and others 1977b). Because seeds There are approximately 30 species of burnet, are readily eaten by rodents (Everett and others 1978b) occurring primarily in Europe and the Middle East and rabbits, little natural increase has occurred on (Hermann 1966). Two species, western burnet and many areas. Where seeds are allowed to mature and Alaskan burnet, are native to Western North America. rodent and rabbit populations are low, reproduction They are locally valued as forage plants (Hermann will take place. Dense stands of small burnet have re- 1966; Hitchcock and others 1961). Selections of mained in some seedings for as long as 25 years. Vege- small burnet from Mediterranean and Middle East- tative propagation can be accomplished by dividing ern countries have been more widely adapted to sites and transplanting the somewhat rhizomatous plants. in the Intermountain region than any of the Western North American species. Small burnet has proven to Uses and Management be well adapted to pinyon-juniper, basin big sagebrush, and mountain big sagebrush communities, and Small burnet forage and seeds are highly preferred to drier exposures in the mountain brush and aspen at all seasons by livestock, big game, rodents, and types. Small burnet will grows well on acid or alkaline upland game birds (Autenrieth and others 1982; soils (Plummer 1977). The species is adapted to sandy Wasser 1982). Birds and rodents also make consider- and clay soils, with silty and loamy soils the most able use of the seeds. Seed caches and resulting plants preferred (Wasser 1982). Small burnet has good win- are common. Small burnet is particularly important ter hardiness and some fire and shade tolerance. in late winter, early spring, and late summer when Introduction of S. magnollii, S. dictyocarpum, and other species provide less green forage. Plants pro- S. muricata have exhibited characteristics similar to duce an abundance of basal leaves that remain green those of S. minor when grown on southwestern Idaho and persist throughout the dry summer and winter rangelands (Shaw and Monsen 1983a). months. Stand vigor and density may be reduced by selective grazing during these periods. Plant Culture Once established, small burnet can compete fairly well with cheatgrass. This forb has done well on Excellent seed production, high seed quality, and mountain brush, pinyon-juniper, and upper sage- ease of seed processing and planting have contributed brush sites. Although it exhibits useful attributes to the widespread use of small burnet. Aerial or hand for the drier sagebrush and desert shrublands, it is broadcasting, drilling, and dribbling have all proven not adapted to areas receiving less than 10 inches to be successful means of seeding. Seed needs to be (25 cm) of moisture. Under favorable circumstances,

458 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats seedlings will establish when planted on dry sites. numerous yellow heads about 0.25 inch (1.0 cm) in However, seedlings often fail if seeded when spring diameter with black-tipped bracts (Welsh and others moisture is low. Mature plants are hardy and com- 1987). Flowering occurs in May to July. petitive and will persist, once established, on areas receiving 8 to 10 inches (20 to 25 cm) of moisture. Ecological Relationships and Distribution Currently, some ecotypes express much better drought tolerance and longevity than most cultivated strains. A native in all Western States, butterweed ground- These selections offer opportunity to expand the use sel (fig. 37) can be found in plant communities ranging of this forb to drier environments. from sagebrush to subalpine. Welsh and others (1987) Seeded areas should not be grazed for at least 2 years describe two varieties with somewhat separate areas of occurrence. Senecio serra var. serra exists in the following seeding. Because plants live from 7 to 12 sagebrush, pinyon-juniper, mountain brush, lodgepole years, every effort should be made to ensure that seed pine, and spruce-fir communities at 6,000 to 10,000 ft crops are not grazed and are allowed to mature every (1,800 to 3,000 m). Senecio serra var. admirabilies 3 to 4 years. grows in ponderosa pine and subalpine meadow com- Small burnet can be seeded, grazed, and maintained munities (Welsh and others 1987). Both varieties are in mixed grass and broadleaf herb plantings. Too early colonizers of disturbed sites and grow on sites often, the species is heavily grazed with livestock in ranging from open, dry slopes with sandy to gravelly the spring and early summer, and later grazed by loam soils to forest openings. Both varieties grow well game animals in the fall and winter periods. Use can in association with wheatgrasses, bromes, wildryes, be expected during any season as the plant retains and a large number of forbs. some green foliage throughout the entire year. Ro- dents, insects, and game animals will forage on the plant whenever it is available. Consequently, livestock grazing must be regulated to compensate for wildlife uses. The plant should be seeded heavy enough to lessen concentrated use and satisfy animal needs. If seeded in mixtures with perennial grasses, the species can diminish in density because it is short- lived. Competitive grasses and heavy grazing can prevent natural seeding. Sufficient plants have persisted on most wildlife plantings where livestock grazing has been controlled.

Varieties and Ecotypes ‘Delar’ small burnet was released in 1979. It is recommended for sites in the Intermountain region that receive at least 12 inches (30 cm) of annual precipitation. It will, however, establish and do fairly well with as little as 10 inches (25 cm) of precipitation. Forage production is excellent, even the first growing season. Under irrigation ‘Delar’ has produced up to 1,050 lb seed per acre (1,177 kg per ha) (Howard 1981). Additional selections are being investigated. There are some selections that do well with as little as 7 inches (18 cm) of annual precipitation, but these strains are not commercially available.

Family Compositae Senecio serra______Butterweed Groundsel This native perennial is a member of the sunflower family. It grows in clumps of several stems 2 to 6 ft (0.6 to 1.8 m) tall and has a woody base. It produces Figure 37—Butterweed groundsel in full bloom.

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Plant Culture Family Compositae Seeds usually mature in September, and can be Solidago canadensis ______collected by hand stripping or by using small paddles to beat them into collection trays. Seeds are attached Canada Goldenrod to a white pappus that should be removed to facilitate A native perennial composite, Canada goldenrod seeding. This can be accomplished by mechanical (fig. 38) produces stems 1 to 6 ft (0.3 to 1.9 m) tall that debearding and screening. Seeds are very small, with grow from creeping rhizomes. Flowering occurs from just under 3.5 million per lb (7.7 million per kg) at 100 July to October. Pale golden-yellow flowers occur percent purity and with the pappus removed. Seed is along only one side of the curved flowering stalk. best broadcast planted unless the pappus is removed, Basal leaves are deciduous or become withered at after which seed can be drilled. Acceptable purity and the time of anthesis (Welsh and others 1987). germination is 50 to 70 percent. This species does well when seeded in mixtures with Ecological Relationships and Distribution other herbs because it has a fairly strong seedling. Fall seeding is preferred. Seed should not be covered more Canada goldenrod occurs from southwestern than 0.13 inch (3 mm). Canada to New Mexico and California in pinyon- juniper, mountain brush, ponderosa pine, and aspen- spruce-fir types. It occurs along riparian areas, in Uses and Management moist places and forest openings, and on dry slopes. It Butterweed groundsel is grazed extensively by sheep, grows on basic, neutral, and slightly acidic soils. cattle, and big game (USDA Forest Service 1937). It greens up early in the spring and retains a large Plant Culture amount of green growth late into the summer. It can Flowering occurs during midsummer, and seeds tolerate considerable grazing, trampling, and drought. mature in October. Generally, an abundance of seeds It persists well in partial shade and frequently re- are produced each year. Seeds are best collected by mains green late in the season. Deer and elk (Kufeld hand stripping or beating. The pappus can be removed 1973; Kufeld and others 1973) make good use of it in from the seed using a debearder. Seeds are then the spring and early summer. Cattle and sheep may separated from the debris with a screening type forage on the entire plant late in the season when the cleaner. The seed pappus is generally removed to leaves are partially dry. Because of its woody base and facilitate seeding. Cleaned seeds can be drilled or stout, fibrous, rhizome system, this species does an broadcast planted. Uncleaned seeds can only be hand exceptionally good job of stabilizing disturbed soil. It broadcast. Seeds should not be covered more than can be used to plant both dry and wet sites, although 0.13 inch (3 mm) deep. Good results can be expected it does not tolerate prolonged flooding. This species can be seeded in disturbed riparian sites where the water level has been lowered by down cutting of the channel. It establishes well from broadcast seeding and can be planted with a grass-forb mixture. Young plants persist well amid competition, but the plants are slow to attain maximum stature. Because it is slow to achieve dominance, it can be seeded with slower developing shrubs and trees without limiting their survival. Young plantings are usually not adversely effected by grazing. Plants normally require 2 to 3 years to fully establish. When planted with a grass mixture, this forb persists equally well and does not require special management. Established stands should be allowed to set seed periodically.

Varieties and Ecotypes There are no releases. Figure 38—Canada goldenrod.

460 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 19 Forbs for Seeding Range and Wildlife Habitats from broadcast seeding on prepared seedbeds and on Ecological Relationships and Distribution disturbed or eroding soils. Once established, plants spread well from creeping rhizomes. Wildland col- This species is a drought-tolerant native forb that lected seed lots are generally highly viable. Only fresh occurs in blackbrush, shadscale, rabbitbrush, sage- seed should be planted because viability declines with brush, and pinyon-juniper communities, and occa- 1 year of storage. This species does well when seeded sionally in the mountain brush zone on sites receiving in mixtures with other herbs because it has fairly 8 to 12 inches (200 to 300 mm) of annual precipitation vigorous seedlings. More successful stands usually (Stevens and others 1985c). It is common on disturbed establish from fall seedings. and burned pinyon-juniper sites (Everett 1982; Plummer and others 1968). It ranges throughout the Uses and Management West from Washington to Arizona (Welsh and others 1987). Canada goldenrod is palatable to livestock and big Gooseberryleaf globemallow prefers full sunlight. It game. Some use is made at most seasons. Plants green exhibits good winter hardiness, grazing tolerance, up early in the spring and remain green longer in the resistant to burning, and considerable drought resis- fall than most associated species. Canada goldenrod tance. Gooseberryleaf globemallow is found in alka- invades disturbed areas and provides an effective line soils and tolerates moderate salinity, but not sodic ground cover. It can be used to control erosion on soils (Pendery and Rumbaugh 1986). extremely disturbed areas and is often used in watershed plantings. This species can be used to beautify Plant Culture recreational areas, summer home sites, administrative areas, and other areas with high aesthetic value. Gooseberryleaf globemallow can be established on It is adapted to semiwet situations and can be used to harsh sites. It is one native forb that has been success- treat riparian disturbances where dry and semiwet fully seeded in blackbrush, shadscale, pinyon-juniper, sites are closely aligned. Canada goldenrod estab- and sagebrush communities and in disturbances with lishes well by transplanting and can be used to provide basic soils (Monsen and Plummer 1978; Plummer immediate ground cover. It persists with heavy tram- 1966, 1977). pling and grazing disturbances. Canada goldenrod usually occurs in aspen and conifer forest communities. It recovers well following burning, logging, and grazing. Established stands should be allowed to set seed periodically. Newly seeded stands should not be grazed for at least 2 years following planting.

Varieties and Ecotypes There are no releases.

Family Malvaceae Sphaeralcea grossulariifolia ______Gooseberryleaf Globemallow There are approximately 200 species of globemallow; about 20 are native to Western North America (Hermann 1966). A member of mallow family, the genus is variable and complex, and its taxonomy has been extensively revised (Jefferies 1972; Kearney 1935; Welsh and others 1987). Gooseberryleaf globemallow (fig. 39) is a densely, gray-green, pubescent perennial. Leaves are gooseberry-like. Floral stalks grow from 1 to 2.5 ft (30 to 76 cm) tall. Plants develop a branched taproot, and a near-surface fibrous root system. Flowers have showy orange to reddish petals (Hitchcock and others 1961). Flowering occurs from May to July, and seed matures unevenly Figure 39—Gooseberryleaf globemallow. from June to August.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 461 Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Seeds can be collected by hand or with mechanical Varieties and Ecotypes strippers. They are easily cleaned. There are just over 500,000 seeds per lb (1.1 million per kg) at 100 percent There are no releases. purity. Seeds should be dusted with an appropriate insecticide to prevent destruction by weevils (Pendery Family Malvaceae and Rumbaugh 1986). Seed viability is, however, maintained for up to 15 years without any significant loss Sphaeralcea coccinea______(Stevens and Jorgensen 1994). Page and others (1966) Scarlet Globemallow reported that low germination results from a combination of low fill and a hard seedcoat impregnated with Scarlet globemallow (fig. 40) is a low-spreading, native nonwetable substances. Mechanical or acid scarifica- perennial with creeping rhizomes (Arnow 1971; McKell tion can increase germination. It can be seeded on and others 1979; Welsh and others 1987). It prefers disturbed and burned pinyon-juniper areas. Because clay soils and can be found on hills and dry plains in the species has fairly strong, competitive seedlings blackbrush, shadscale-greasewood, sagebrush, pinyon- and a moderate growth rate, it can be seeded in juniper, mountain brush, and ponderosa pine commu- mixtures with other species. Fall and winter seeding is nities from Southern Canada to Texas and Arizona recommended (Monsen and Plummer 1978). Seeds (Hermann 1966; Welsh and others 1987). can be drill seeded and should be planted at depths of Scarlet globemallow exhibits considerable drought 0.25 inch (6 mm). Seeds may lie in the soil for years and resistance, and establishes especially well on dis- germinate when conditions are suitable (Plummer turbed sites. This species can be successfully seeded or and others 1968). This species can be successfully transplanted in blackbrush, shadscale, black grease- transplanted and will reproduce from stem cuttings wood, sagebrush, and pinyon-juniper types. It is par- (Everett 1982). ticularly useful for planting on disturbed sites, as it is drought tolerant and can spread by rhizomes. It is Uses and Management well suited to adapted sites where wildfires frequently occur. Forage value of gooseberryleaf globemallow is rated Palatability for livestock and big game is rated from as good for sheep and cattle (Hermann 1966; Wasser poor to good (Hermann 1966; Kufeld 1973; Kufeld and 1982). Preference or palatability is rated as none to others 1973; McKell and others 1979; Wasser 1982). fair for cattle and sheep (Hermann 1966), and fair to excellent for antelope (Hancock 1966; Smith and Beale 1980), elk, and deer (Kufeld 1973; Kufeld and others Varieties and Ecotypes 1973; Urness and McCullock 1973). Antelope espe- There is one released germplasm, ‘ARS 2936’. cially prefer this herb when it is flowering (Pendery and Rumbaugh 1986). Protein content during winter and early spring has been found to be high, ranging from 10 to 20 percent (Pendery and Rumbaugh 1986; Urness and McCullock 1973). Gooseberryleaf globemallow greens up early in spring and will green up again in autumn after fall storms. This species establishes well on disturbed sites from seed or from transplanting. Because of its drought resistance, continuous, colorful flowering, and ease of establishment, this species has been recommended for use in dryland ornamental landscaping (Natural Veg- etation Committee 1973). Gooseberry globemallow is quite competitive and can be used to suppress cheatgrass and other annuals. It is one of only a few native herbs that can be seeded on low elevation Wyoming big sagebrush and shadscale sites. Following germination gooseberry globemallow produces strong and persistent seedlings. It has a deep taproot. Plants are tolerant of grazing, but can be weakened by close grazing in late spring. Drought tolerance is excellent, but plants should not be grazed following extended periods of drought. Plant density tends to fluctuate from year to year. Figure 40—Scarlet globemallow.

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Family Fabaceae Family Fabaceae Trifolium ______Trifolium fragiferum ______Clovers Strawberry Clover Taylor (1985) reported that about 250 species of true Strawberry clover was introduced to the West from clovers occur throughout the world. All species possess Eurasia and the Mediterranean region (Forde and the typical papilionaceous legume flower with 10 sta- others 1981). It is not known when it was introduced mens. Leaves usually consist of three leaflets. Most into the United States, but specimens were collected have simple taproots; some produce stolons or rhi- in 1878 in Pennsylvania. zomes. About one-third are perennials and the others Strawberry clover is a low-growing stoloniferous are annuals. About one-third are self-pollinated; the perennial used primarily as a pasture legume others are cross-pollinated and require bees for polli- (Townsend 1985). Stems are decumbent to creeping nation (Taylor and others 1980). All species are able and root at the nodes. Plants are less than 10 inches to fix nitrogen if nodulated with strains of Rhizobium (25 cm) tall. White to pink flower heads are borne at (Allen and Allen 1981). the ends of each seed stalk (Gillett 1985; Harrington Within the contiguous United States, clovers are 1964; Hermann 1966). The flower and seed head most important as range plants from the Rocky Moun- resemble a strawberry—hence the name. Plants are tains west to the Pacific Coast (Crampton 1985). pubescent or glabrous. The leaves are congested on Approximately 65 species occur in the Western States. Most species, particularly many annuals, are abun- auxiliary peduncles that are curved and ascending dant in the central valley of California (Crampton (Bendixen and others 1960). 1985; Küchler 1977). Although native perennial clovers are found abun- Ecological Relationships and Distribution dantly throughout the mountains, meadows and wet- Strawberry clover has spread or been seeded through- lands of the Intermountain area, and to a lesser extent in the drier sagebrush habitats (Crampton 1985), few out North America and has become naturalized in natives have been utilized in range revegetation ef- some locations (Townsend 1985). It is noted for its forts. Most clovers of agricultural interest were intro- ability to grow on wet saline or alkaline soils and has duced from Europe. There are about 15 species that demonstrated adaptability to wet meadows, streams, are used in North America (Hermann 1953; Hollowell and seeps. Bendixen and Peterson (1962) report that 1960). Most are adapted to temperate climates and are strawberry clover withstands flooding because the used to improve irrigated or native meadow pastures. stolons exhibit a tropic response that causes the tips to Because they are legumes, clovers are particularly be elevated above the water level. Light, oxygen defi- useful in fixing atmospheric nitrogen and enhancing ciency, and atmospheric conditions also influence the soil fertility. A cool, moist climate is required for the tropic response. best growth (Taylor 1985). Spring and fall periods are Strawberry clover is not hindered by wet-saline favorable for adequate growth in the Intermountain conditions; consequently, it is very useful on many area.Various species demonstrate usefulness on a seeps and salty sites at a wide range of elevations. variety of sites, and future trials will undoubtedly Seedlings are not suppressed by high osmotic condi- result in more widespread use of different clovers. tions that depress the growth of white clover (George Most clovers are quite palatable, shade tolerant, and and Williams 1964). tolerant of flooding for short periods. Consequently, they are useful for seeding semiwet meadows, and as Plant Culture understorys in conifer forests and aspen stands. All species attract use by wildlife and can be planted to Strawberry clover can be drill or broadcast seeded supply forage for extended periods. Most species demon a prepared seedbed. Seed coverage should be shal- onstrate good regrowth and, thus, withstand heavy low, less than 0.25 inch (6.4 mm). When seeding irri- grazing. gated pastures, a planting depth of about 0.50 inch Seeds are small and include a high percentage of (13 mm) is recommended (Townsend 1985). Seeds are hard seeds. Seeds should be treated with an inoculum small with about 290,000 seeds per lb (650,000 per kg) to assure establishment. at 100 percent purity. They require planting into a firm seedbed. Evers (1982) found that drill seeding

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 463 Chapter 19 Forbs for Seeding Range and Wildlife Habitats produced only slightly higher yields than broadcast- spring or fall plantings. Drilling or broadcast seeding ing. Seeding rates vary depending on seeding meth- on a firm seedbed is recommended. Planting with ods, but acceptable stands are achieved with 6 to 8 lb grasses to improve forage productivity is a common per acre (6.7 to 9.0 kg per ha) when seeded alone, and practice (Grable and others 1965). 2 to 4 lb per acre (2.2 to 4.5 kg per ha) when planted in Seeding in mixtures at a rate of 2 to 4 lb per acre a grass mixture. (2.2 to 4.5 kg per ha) produces excellent stands. There Strawberry clover seedlings are easily established are approximately 68,000 seeds per lb (150,000 per kg). and grow rapidly. The plant establishes well with Seeds have a hard seedcoat, but this does not prevent minimal seedbed preparation. Successful stands es- good establishment even from spring seedings. tablish from broadcast seeding and natural spread. No cultivars have been developed for rangeland This clover does well when seeded in mixtures with conditions in the United States, but two Canadian grasses. When planted in mixtures under irrigation, cultivars, ‘Aurora’ and ‘Dawn’, have been developed herbage production can be increased and better dis- for pasture plantings. tributed throughout the growing season (Peterson and others 1962). Family Fabaceae Uses and Management Trifolium pratense ______Forage and seeds of strawberry clover are used Red Clover extensively by livestock, big and small game, and upland game birds. Red clover is perhaps the most widely used pasture Strawberry clover is especially well suited to close clover in the United States. It is generally grown in and continuous grazing (Raguse and other 1971), and mixtures with grasses for intense pasture manage- when seeded on most rangeland sites, tends to attract ment. It is also grown for hay or silage production, or heavy use. In vitro digestibility compares favorably as a soil improvement crop. Plants produce nitrogen with white clover, red clover, and alfalfa (Reed and that favors growth of companion species and im- others 1980). proves soil conditions (Smith and others 1985). This forb can be successfully seeded onto depleted Red clover can be used as a short-term perennial or riparian habitats, degraded meadows, and inland winter annual to furnish immediate forage and im- saltgrass areas. It can be used to stabilize streambanks, prove soil fertility. It requires 20 or more inches (51 erodible soils, and related disturbances. It is particu- cm) of annual precipitation, but it is well suited to cut larly useful for seeding disturbances on saline soils over timber lands and moderately acid soils and sites subjected to frequent periods of flooding. (Hafenrichter and others 1968). It can be used to improve meadows and degraded riparian sites. Varieties and Ecotypes Hafenrichter and others (1968) reported that the commercial varieties—‘Kenland’, ‘Pennscott’, ‘Dollard’, No selection or cultivar has been developed for and ‘Lakeland’ are well adapted to conservation plant- rangeland uses, although various cultivars have been ings in the Northwest. None have been extensively developed for pasture seedings in the United States evaluated for range sites in the Intermountain area. (Peterson and others 1962). Family Fabaceae Family Fabaceae Trifolium repens ______Trifolium hybridum ______White Clover Alsike Clover White clover is a shallow rooted perennial with Alsike clover is a native of northern Europe that was creeping stems that root at the joints. This introduced introduced into the United States in about 1839 European species has become naturalized in North (Townsend 1985). It was thought to be a hybrid of red America in fairly moist areas on medium and high and white clover—hence its species name “hybridum”. elevation mountain ranges. White clover is winter It is well adapted to high elevations within the Inter- hardy and can withstand more extreme temperatures mountain region where conditions are cool and moist. than either red or alsike clover (USDA Forest Service It is a short-lived perennial, but can be managed to 1937). Livestock and wildlife make good use of this reseed and maintain itself. Alsike clover is an excel- species. Areas of adaptation include sites with fertile lent hay and pasture species (Townsend 1985), but is and well-drained soils that are moist throughout the also useful as a range forage species. It is adapted to growing season, and along streams. It has good soil sites that are too wet or acidic for red clover or alfalfa stabilization characteristics. White clover is generally (Townsend 1985). Plants establish well from either seeded in a mixture with perennial grasses.

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Family Compositae Viguiera multiflora var. multiflora __ Showy Goldeneye There are about 150 species of Viguiera. Most species occur in the new world (USDA Forest Service 1937). Two varieties of goldeneye, showy goldeneye and Nevada goldeneye, occur within the Intermoun- tain region (Welsh and others 1987). Showy goldeneye (fig. 41), a member of the sunflower or compositae family, is a perennial native Figure 42—Showy goldeneye in an aspen opening. forb growing from a short, branched, woody taproot. It is usually 1 to 3 ft (30 to 91 cm) high with leaves commonly more than 0.25 inch (6 mm) wide and 0.50 to 0.75 inch (12 to 19 mm) long. Two or more showy, yellow, sunflower-like blossoms (heads) are borne on each plant. Flowering begins in July. Seed matures in Ecological Relationships and Distribution late August and September. Basal leaves elongate and green up shortly following snowmelt (Welsh and oth- Showy goldeneye is a very attractive perennial, that ers 1987). occurs throughout the Intermountain West at elevations between 3,500 to 11,000 ft (1,100 to 3,400 m) (Stevens and others 1985c). Showy goldeneye can be found from moderately moist habitats to dry, open, rocky slopes in sagebrush-grass, pinyon-juniper, mountain brush, aspen (fig. 42), spruce-fir, subalpine, and often in riparian communities (Welsh and others 1987). This species is not restricted to full sunlight, but can be found in partial shade and in densely wooded areas (Arnow 1971; Plummer and others 1955). Showy goldeneye can be found on a wide variety of soils ranging from heavy clays to gravel (USDA Forest Service 1937), and from acidic to basic conditions (Plummer 1977). It establishes quickly on disturbed sites, has a rapidly developing seedling, and competes well with annuals and perennials.

Plant Culture Each showy goldeneye plant can support numerous, golden yellow, sunflower-like heads that generally produce an abundance of seeds. Seeds are collected by hand, mechanical beating, or stripping. Viability of wildland lots is only fair. Seed can be stored up to 7 years without any major loss in viability (Stevens and Jorgensen 1994). There are just over 1,000,000 seeds per lb (2.2 million per kg) at 100 percent purity. Showy goldeneye develops strong seedlings. Plants establish quickly, and can spread from seed. Showy goldeneye will invade disturbed sites and may quickly become the dominant species. Seeds can be broadcast or drilled with equal success. Seeds should not be planted more than 0.25 inch (6 mm) deep. Seeding is best accomplished in the fall or winter. The species does well when planted as a component of a mixture Figure 41—Showy goldeneye. (Stevens and others 1981b).

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Principal Areas of Use Family Compositae Showy goldeneye is an early spring-greening forb Viguiera multiflora var. that is sought by big game (Kufeld 1973) and livestock. nevadensis______Foliage and flower heads are readily consumed by cattle, sheep, deer, and elk. Birds also make consider- Nevada Goldeneye able use of the seeds. Strong seedlings and a rapid rate of growth enable Compared to showy goldeneye, Nevada goldeneye showy goldeneye to be seeded in conjunction with has narrower leaves (usually less than 5 mm wide) and other species. These features also contribute to its can be found on drier sites in southwestern Utah, establishment and spread into annual communities. Nevada, and California. It occurs with pinyon and Profuse flowering of this species makes it a prime juniper, Wyoming and basin big sagebrush, black candidate for summer home sites, campgrounds, ad- sage, and various saltbushes. Plummer (1966) recom- ministration sites, and other areas with high aesthetic mends it as a candidate species to be used in seeding values. This species has potential as an erosion control areas in the salt desert shrublands. Coles (1982) species on disturbed and burned sites. reported that Nevada goldeneye rapidly invaded Wyo- Newly seeded areas should not be grazed for at least ming and basin big sagebrush burns in southwestern 2 years following planting. Established stands should Utah. He further reported that mule deer tend to seek be allowed to set seed every 3 to 4 years. out and concentrate on areas with high densities of Nevada showy goldeneye. The species invades road- Varieties and Ecotypes sides and other disturbed areas. Seed production and seeding procedures are similar to those described for There are no releases. showy goldeneye.

466 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 E. Durant McArthur Stephen B. Monsen Chapter 20

Chenopod Shrubs

Chenopod plants (Family Chenopodiaceae) are distributed worldwide but are especially prominent in some wet and dry saline or alkaline situations. Chenopods are both herbaceous and woody. The relative proportions of life-forms in the family is demonstrated by data from the important center of chenopod diversity in south-central Asia, where n = 341 species: 76 percent are herbaceous (mostly annual), 23 percent are shrubs or subshrubs, and 1 percent is arborescent (McArthur and Sanderson 1984; Shishkin 1936). Chenopod shrubs grow over wide expanses of Intermountain rangelands as well as on large saline and alkaline tracts on all continents, except Antarctica. In the Inter- mountain area, seven genera (table 1) with 28 species make important contributions to the landscapes or to revegetation needs. As a group the chenopod shrubs exhibit the ability to grow in low fertility, salt-bearing soils as well as on more favorable sites. Their presence is essential for maintaining a stable soil in many xeric environments where soil is too salty or dry for most other classes of plants to live (Blauer and others 1976; Goodall 1982; Sanderson and Stutz 1994b; Wilkins and Klopatek 1984). Cheno- pod shrubs grow well in high concentrations of calcium and potassium salts and can endure considerable concentrations of

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Chapter Contents by Species

Atriplex canescens (fourwing saltbush)...... 471 Atriplex confertifolia (shadscale) ...... 476 Atriplex corrugata (mat saltbush) ...... 479 Atriplex cuneata (Castle Valley clover or cuneate saltbush) ...... 480 Atriplex gardneri (gardner saltbush) ...... 481 Gardner saltbush relatives ...... 481 Atriplex buxifolia ...... 481 Atriplex falcata ...... 481 Atriplex tridentata ...... 481 Other Saltbushes ...... 482 Atriplex lentiformis ...... 482 Atriplex obovata ...... 482 Atriplex polycarpa ...... 482 Atriplex hymenelytra ...... 482 Ceratoides lanata (winterfat) ...... 484 Grayia brandegei (spineless hopsage) ...... 486 Grayia spinosa (spiny hopsage)...... 487 Kochia americana (gray molly)...... 488 Kochia prostrata (forage kochia) ...... 489 Sarcobatus vermiculatus (black greasewood) ...... 490 Other chenopod shrubs...... 491 Allenrolfea occidentalis (iodine bush) ...... 491 Camphorosoma monspeliaca (Mediterranean camphorfume) ...... 491 Suaeda torreyana (desert sumpbush, desert seepweed, or desert blite) ...... 491

sodium salts (Gates and others 1956; McNulty 1969; reported consistent morphological differences among Moore and others 1972). Certain ones grow through a the chromosome races of fourwing saltbush. Even much wider range of alkalinity than others (table 2). though genetic factors are important, moisture does, A few tolerate such high salt concentrations that they of course, markedly affect luxuriance and growth. are the only plant life. As a group they are especially Wallace and others (1974) reported that 40 percent of suited for planting on depleted salt desert shrub type the biomass of fourwing saltbush is in the root system. range areas as well as on construction scars, mine Shrubby chenopods have a chromosome number spoils, and roadcuts (McArthur and others 1978b; based on x = 9. Polyploids occur in most genera Plummer 1977). (Blauer and others 1976; McArthur and Sanderson Within species, large differences exist between 1984). They are wind pollinated, occur in sympatric ecotypes in morphological and physiological traits. distributions, are monoecious, dioecious, or even The smaller ecotypes are usually from dry, severe trioecious—factors that lead to common interspecific sites, whereas the larger ones are from more favorable hybridization (McArthur 1989). Woody chenopods areas. Size of the shrubs is, to a large extent, geneti- show remarkable capability to hybridize both natu- cally controlled as demonstrated by maintenance of rally (Sanderson and Stutz 1994b; Stutz 1978, 1984) ecotypic form in accessions grown together in uniform and under controlled conditions (Blauer and others gardens (McArthur and others 1983a). Furthermore, 1976; Drobnick and Plummer 1966; Stutz 1982). Stutz and Sanderson (1983) have shown in shad- Most of the hybridization occurs within Atriplex, scale that the smallest ecotypes growing on the most although some possible intergeneric hybrids have severe sites are higher polyploids, whereas lower been reported (Blauer and others 1976). ploidy levels occur on more favorable mesic sites. Hybridization allows opportunity to create new The gigas form of fourwing saltbush reported by gene combinations to fill new niches—for example, Stutz and others (1975) is a diploid. Dunford (1985) those in newly created mine spoils. Stutz (1982),

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Table 1—Shrubby chenopods in Western America useful or Ferguson and Frischknecht (1985), and Stutz and potentially useful for restoration and revegetation Estrada (1995) found that several hybrid combina- (adapted and expanded from McArthur and others tions within Atriplex performed well on mine spoils in 1978b). Utah and New Mexico. Blauer and others (1976), Scientific name Common name Carlson (1984), Stutz (1983, 1984), and Stutz and Carlson (1985) showed that there is ample opportu- Allenrolfea occidentalis Iodine bush or inkbush nity to: (1) make selections from natural populations, Atriplex aptera Wingless saltbush (2) locate natural hybrids, and (3) make artificial Atriplex bonnevillensis Bonneville saltbush Atriplex canescens Fourwing saltbush hybrids. All three plant classes can be useful in provid- Atriplex confertifolia Shadscale ing plant materials to rehabilitate harsh disturbed Atriplex corrugata Mat saltbush landscapes. Sanderson and colleagues (Sanderson Atriplex cuneata Castle Valley clover saltbush and Stutz 1984, 1994a; Sanderson and others 1988, or cuneate saltbush 1990) have identified flavonoid compounds that are Atriplex falcata Falcate saltbush useful genetic markers in population biology and Atriplex gardneri Gardner saltbush hybridization studies. Atriplex hymenelytra Desert holly saltbush Woody chenopods show remarkable ability to grow Atriplex lentiformis Big saltbush or quailbush through a wide climatic variation. Some grow in the Atriplex navajoensis Navajo saltbush coldest parts of the northern desert shrublands and Atriplex obovata Broadscale saltbush adjacent grasslands to warmer desert ranges in the Atriplex polycarpa Cattle saltbush or allscale saltbush Southwestern United States and Mexico (fig. 1). Atriplex robusta Robust saltbush Three examples are Gardner saltbush, black grease- Atriplex semibaccataa Australian saltbush wood, and winterfat. Several chenopods readily estab- Atriplex tridentata Trident saltbush lish, grow, and produce in areas of severe aridity or Camphosoma monspeliacaa Mediterranean where the average annual precipitation may be less camphorfume than 5 inches (12.7 cm). Important among these are Ceratoides lanata Winterfat winterfat, shadscale, and falcate saltbush. However, a Ceratoides latens Pamirian winterfat they can also quickly take advantage of three to four Grayia brandegei Spineless hopsage times this precipitation to make luxuriant growth. Grayia spinosa Spiny hopsage While the shrubby chenopods are most characteris- Kochia americana Gray molly tic of the salt desert shrub type in Western North Kochia prostrataa Forage kochia Sarcobatus vermiculatus Black greasewood America, some grow well in limey soils on mountain Sarcobatus baileyi Bailey greasewood ranges in association with ponderosa pine, Utah Suaeda suffrutescens Desert sumpbush juniper, pinyon pine, Saskatoon serviceberry, and Zuckia arizonica Arizona zuckia Utah serviceberry. Winterfat, fourwing saltbush, and aIntroduced species. spiny hopsage grow best in these areas. However, on

Table 2—Physical dimensions and tolerance to alkalinity of some chenopod shrub taxa (from McArthur and others 1978b).

Adaptation to alkalinity Species Height/crown Optimum Range Inches (cm) Atriplex canescens 12-78/12-178 (Fourwing saltbush) (30-200/30-450) Low Low-medium Atriplex confertifolia 6-47/6-30 (Shadscale) (15-120/15-75) High Low-high Atriplex corrugata 1-8/8-40 (Mat saltbush) (2-20/20-100) Medium Medium-high Grayia spinosa 12-47/12-70 (Spiny hopsage) (30-120/30-180) Low Low Ceratoides lanata 2-47/6-30 (Winterfat) (6-120/15-75) Low Low-medium Allenrolfea occidentalis 6-47/12-70 (Iodine bush) (15-120/30-180) High High Sarcobatus vermiculatus 40-118/40-78 (Black greasewood) (100-300/100-200) Medium Low-high

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 469 Chapter 20 Chenopod Shrubs

before germination. This may provide some survival value for the species because it ensures that some seeds remain in the ground to germinate and establish British when temperature and moisture are most favorable.

Columbia Alberta Apparently, saltbush seeds are not highly palatable Saskatchewan to small rodents (Everett and others 1978) so they are retained in the soil. Some buildup occurs in the ground of fruits that have a tendency for delayed Wash . germination. Consequently, in favorable years many

N. D Montana ak. seedlings become established. Young seedlings, how- Oreg. ever, are particularly susceptible to rodent depreda- Idaho S. Dak. tion (Nord and Green 1977; Young and others 1984e). Wy oming Nevada Shrubby chenopods express a number of traits

Utah Neb. that aid in artificial revegetation. The appendages Calif. Colo. attached to the utricles can be removed from most Kan. species. Except for winterfat, all seeds can be cleaned A rizona New Mexico Okla. to a form that is easily handled and planted with conventional equipment (chapter 19). Cleaned seeds are of a size and shape that different seeding rates Texas can be employed. Within a specific collection, seed germination features are uniform. However, seed germination is not so specific to a particular set of

Mexico Shrubby chenopods climatic conditions that all seeds germinate at any Ceratoides lanata one time. An extended period of germination often Atriplex gardneri complex assures the survival of a satisfactory number of Sarcobatus vermiculatus plants. However, seed germination of winterfat is usually confined to a short duration. Seedlings and young plants are quite vigorous and are able to compete well with herbs (Giunta and others 1975). The growth rate of nearly all shrubs is excellent. Figure 1—Distribution of shrubby chenopods Even under adverse conditions plants grow remark- in general and winterfat, greasewood, and the ably well, often exceeding most herbs. Plants are well Gardner saltbush complex in particular. Distri- adapted to infertile soils and can be used to restore butions from Hall and Clements (1923), Branson vegetation throughout variable, but harsh, situa- (1966), and McArthur and others (1978b). tions. Young plantings withstand heavy browsing by game, livestock, and rodents. Individual plants normally reach maturity within 3 to 5 years, and natural spread can be expected thereafter. Once established, highly alkaline extrusions in these higher elevations young plants persist under adverse climatic condi- are precipitation areas on mountain ranges where tions. In addition, established plants provide high shrubby chenopods may grow to the virtual exclusion yields even when grown with a dense understory. of other species (McArthur and others 1978b). Chenopod shrubs frequently grow with a sparse All shrubby chenopods bear a carpellate, loculed understory but persist and grow well when seeded one-celled fruit known as a utricle, within which is a with numerous other plants. These shrubs normally single seed. Actual seeds are small, usually no larger occur throughout some principal sites or vegetative than an alfalfa seed. The utricle wall may be quite conditions important to both livestock and wildlife. thin and easily removed from the seed, as in winter- The plants can be increased in density or seeded with fat; or may be hard, as found in shadscale and some a combination of herbaceous plants to improve forage accessions of fourwing saltbush. Production of fruits or habitat. Many other shrubs exist within specific varies widely among plants and from season to season site conditions and do not grow well when moved to as does percentage fill (Crofts 1977; Gamrath 1972; adjacent locations. Some ecotypes of many chenopod McArthur and others 1978b; Monsen and McArthur shrubs also exhibit specific site adaptation require- 1985; Springfield 1970b). To a considerable degree ments, but others can be extended to additional sites this relates to how favorable the growing season may (see McArthur and others 1983a). be. Unless seed coats are restrictive, germination Most shrubby chenopods are highly nutritious. They takes place quickly in a favorable environment of have higher protein content than most plants, par- temperature and moisture. Utricles having indurated ticularly in the winter (Chatterton and others 1971; walls, such as in some ecotypes of fourwing saltbush Esplin and others 1937; Tiedemann and others 1984a; and shadscale, may lay a year or more in the ground Welch and Monsen 1981). On several species and

470 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 20 Chenopod Shrubs accessions, leaves persist and remain green well because of the white, scurfy vestiture. The linear to through winter. These are preferred by grazing ani- oblanceolate or spatulate evergreen leaves are 1 to 4 mals during the winter and provide an important cm long and 2 to 6 mm wide. Fourwing saltbush is source of protein, which is vital to the well being of usually dioecious with a low frequency of monoecy. browsing animals on desert ranges of the West. Pistillate (female) flowers are small and inconspicu- Without doubt, the shrubby chenopods have an ous with no flower parts except pistils. Each pistil is important future for development of cover and forage enclosed by a pair of small bracts that are united along on the generally basic soils of arid ranges world- their edges to form winglike expansions. In addition, wide. They are widely used in land rehabilitation each bract of the pair has a wing down the middle; so programs (Aldon and Oakes 1982; Plummer 1984; the utricle (fruit) has, at maturity, a varying promi- Tiedemann and others 1984b). nence of four wings (fig. 3, 4). The yellow to red to Recently, Nechaeva and others (1977), Rumbaugh brown staminate flowers are more or less naked an- and others (1981, 1982), and Ostyina and others (1984) thers (only a few bracts are present) borne in glomer- showed that most shrubs, especially some chenopods, ules 2 to 3 mm wide (Blauer and others 1976; McArthur provide complementary forage to grasses in mixed and others 1978c). Sex expression in fourwing salt- stands on semiarid ranges. Forage quality is enhanced bush is apparently a dioecious system for the only in the mixed stands and the effective season of grazing diploid population known in Utah (Freeman and is expanded. In addition, the forage resource is made McArthur 1989; McArthur and Freeman 1982; Stutz less susceptible to attack from insects and diseases and others 1975). Barrow (1987) has shown that than are monocultures (Moore and others 1982). diploid New Mexico and Texas populations may con- Nord and Green (1977) identified several chenopod tain rare monoecious plants. At the higher and more shrubs with potential to reduce the hazards of fire in common (tetraploid, hexaploid) ploidy levels, fourwing the California chaparral. Among these were North American fourwing saltbush, Castle Valley clover saltbush, Gardner saltbush, allscale saltbush, and three Australian saltbush species. From 1985 through 1988 a significant die-off of chenopod (and some other families) shrubs occurred A in the Intermountain area (McArthur and others 1990b; Nelson and others 1989; Wallace and Nelson 1990). This die-off was particularly serious for shadscale. The die-off was not unique in time, but it was large in scale; and recovery will be complicated by the recent large-scale invasion of exotic annuals into chenopod shrublands. Important characteristics of a number of chenopod shrubs follow. Seeding recommendation for principal vegetative types and conditions are discussed in chapter 17. Chenopod shrubs adapted to these vegetative types and conditions are included in the seeding rec- B ommendations. Seed characteristics are found in chapters 24 and 26. Included in the remainder of this chapter are a brief species description, ecological relationship, distribution, culture requirements, use, improved varieties, and management of each of the species appearing in the chapter contents list.

Atriplex canescens ______Fourwing saltbush Figure 2—Fourwing saltbush is an upright Fourwing saltbush is an upright shrub (fig. 2). shrub. (A) A large plant along the Snake River, Its height may vary from 1 ft (0.3 m) to 6.7 ft (2.04 m), Payette County, ID, opposite Nyssa, OR. (B) depending on site conditions and genotype. It branches A ‘Rincon’ fourwing saltbush orchard at the freely from the base. Its mature branches are quite Upper Colorado Environmental Plant Center, brittle. Both young leaves and twigs are gray-green Meeker, CO.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 471 Chapter 20 Chenopod Shrubs

AB C

Figure 3—Flowers and fruits of fourwing saltbush (A) A pistillate (female) plant in fruit (McArthur 722, seeded near Mountain Home, Elmore County, ID, from seed collected near Panaca, Lincoln County, NV; 0.65x). (B) A staminate (male) plant in flower (McArthur 378, St. George, Washington County, UT; 0.87x). (C) A monoecious plant (McArthur 723, Payette County, ID; 0.56x).

Figure 4—Fruits (utricles) of various chenopod species. (A) Fourwing saltbush. (B) Shadscale. (C) Mat saltbush. (D) Castle Valley clover saltbush. (E) Trident saltbush. (F) Broadscale saltbush. (G) Hybrid of fourwing saltbush x shadscale. (H) Four- wing saltbush x trident saltbush. (I) Winterfat. (J) Spineless hopsage. (K) Spiny hopsage. (L) Gray molly. (M) For- age kochia. (N) Black greasewood. (After Blauer and others 1976).

472 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 20 Chenopod Shrubs saltbush is trioecious (three sexual states). In the seed orchards that combine desired vegetative and trioecious condition, some plants are lifetime constant floral traits together for seed production (Briggs 1984; females, others lifetime constant males, and some McArthur and others 1978c; Noller and others 1984; plants are apparently able to avoid stressful situa- Van Epps and Benson 1979). Such orchards may not tions by switching away from the physiologically tax- only provide superior seed but may also provide in- ing female state (Barrow 1987; Freeman and McArthur come from marginal agricultural land. 1989; Freeman and others 1984; McArthur 1977; Most wildland stands of fourwing saltbush produce McArthur and Freeman 1982; McArthur and others a harvestable seed crop at least 3 out of every 5 years. 1992; Noller and others 1984). Gender differences Production and seed quality vary among sites, but appear to be adaptive and allow fuller exploitation of fourwing saltbush is an abundant seed producer. Seed habitats (Freeman and others 1993, 1997; Pendleton is collected by hand stripping or beating, although and others 1992). Dr. Daniel J. Fairbanks and his some vacuum and reel-type harvestors have been students at Brigham Young University have made modified for this purpose (Monsen 1978; Monsen and progress toward understanding the molecular genet- others 1985b; Van Epps 1978). Seed is reasonably easy ics basis of sex expression in woody Atriplex (Freeman to harvest and process. Consequently, seed costs are and others 1993, 1997; Randell 1997; Ruas 1997). not excessive. Seed is harvested in late fall and early winter, and seed can be cleaned immediately after Ecological Relationships and Distribution collection. Various commercial growers have recently established seed orchards of fourwing saltbush. The species grows in a variety of soil types from the Seed yields from these centers appear to be more Great Plains to Pacific Coast ranges and from reliable than wildland stands. Seed quality differ- Canada to Mexico (fig. 1); it occurs from elevations ences may be better as well (Briggs 1984; McArthur below sea level to about 8,000 ft (2,400 m). This and others 1978c; Noller and others 1984; Van Epps shrub is well suited to deep, well-drained sandy soil, and Benson 1979). sand dunes, gravelly washes, mesas, ridges, and The wings are removed from the utricles using slopes; vigorous plants have been found in heavy hammermills, flail-type beaters, Dybvigs or cylinder clays. Ecotypes of the species may grow in widely different soil types and plant communities (Dunford grinders (chapter 24). If correctly operated, these 1985; McArthur and others 1978b; Richardson and cleaners can be used to fracture the utricle to aid seed McKell 1980). Fourwing saltbush is frequently found germination. Removal of the wings greatly reduces intermixed with black greasewood, shadscale, Castle the bulk and allows the utricles to be planted with Valley clover saltbush, basin big sagebrush, and occa- most seeding equipment. sionally black sagebrush and pinyon-juniper. In many Only about 50 percent of the utricles contain a full Western regions it is the main shrub over extensive viable seed. Seed “fill” can be increased by cultiva- grassland areas. Important grasses on these tracts tion and culture of nursery plantings. Seed fill from are galleta grass and blue grama. It is not unusual to wildland collections may vary from a low of 10 to find it growing in association with spring-growing 20 percent to as high as 80 percent (Crofts 1977; grasses such as bluebunch wheatgrass and Sandberg Gamrath 1972; Gerard 1978; Springfield 1970b). bluegrass. Seeds should be carefully inspected prior to harvest to ensure the acquisition of high quality seed. Accept- Plant Culture able fill is about 40 percent. Seeds can be stored for a number of years (up to 7) in an open warehouse This species may be propagated by direct seeding, without loss of viability. Even after 15 years storage, transplanting nursery or container stock, or by rooted viability is acceptable (Stevens and others 1981a). stem cuttings (Hennessey 1985; Kay and others 1977b; Seeds of fourwing saltbush normally require 20 to McArthur and others 1978c; Richardson and others 30 days of stratification to initiate and assure uniform 1979; nine papers in the revegetation section of germination. Thick impermeable utricles often re- Tiedemann and others 1984b). Shaw and Monsen quire much longer periods before water and oxygen (1984) give information on growing fourwing saltbush can gain entry. Springfield (1966) and Potter and and other chenopod shrubs in a nursery setting to others (1986) found temperatures required for ger- provide suitable bare root transplant stock. Ferguson mination varied among ecotypes, but the optimum (1980) recommended a growth medium of 50 percent range was 15 to 18 ∞C (59.0 to 64.4 ∞F). Potter and sphagnum peat moss, 30 percent arcillite aggregate, others (1986) reported seeds are able to germinate at and 20 percent vermiculite for growing Bonneville low media osmotic potential if optimum tempera- saltbush and other plant species native to semiarid tures are maintained. However, ecotypes differed in alkaline soils. Use has been made of rooted stem their tolerance to moisture stress. Seeds germinate cuttings (McArthur and others 1984a) to establish early in the spring, and if fall seeded, high mortality

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 473 Chapter 20 Chenopod Shrubs to spring frost can occur. Consequently, spring Uses and Management plantings are sometimes recommended. However, seeds must be planted early enough to facilitate Fourwing saltbush is one of the most valuable forage stratification. If this is not possible or if seeds with shrubs in arid rangelands because of its abundance, impermeable seedcoats or long-term stratification accessibility, palatability, size, evergreen habit, nutri- period requirements are to be planted, fall seeding tive value, rate of growth, and large volume of foliage should be conducted. Where the climate permits, fall (McArthur and others 1978; Petersen and others 1987). or winter planting is preferred. Its leaves, stems, and utricles provide browse in all Seed should not be covered more than 0.5 inch seasons. Sanderson and others (1987) have shown (1.2 cm) deep. Broadcast seeding followed by chaining that saponin content may be important in the palat- has produced excellent stands. Satisfactory stands ability of fourwing saltbush. In addition to providing have also developed from drill and interseeding. Plants forage and cover, this species is one of the most can also establish when seeded in separate rows with important shrubs for use in rehabilitation of depleted herbs using a conventional rangeland drill. However, rangelands and in soil stabilization projects in West- fourwing saltbush seedlings are not able to compete ern desert areas (Blaisdell and Holmgren 1984; Blauer with herbs if drilled together in a single row. Separat- and others 1976; Plummer 1984; Plummer and others ing seeds of different species and reducing herb seed- 1966b). Stutz (1982) and Stutz and Carlson (1985) ling density by broadcast seeding is beneficial to shrub have shown that fourwing saltbush hybridized with establishment. Most satisfactory stands have been other saltbush species can be effective in an in situ achieved when fourwing saltbush was seeded at 1 to selection process in revegetation efforts. Many lines of 2 lb PLS/acre (1.2 to 2.2 kg/ha) or between 2 to 4 lb potentially adaptive plants are established under this (2.2 to 4.5 kg) bulk or cleaned utricles. Between 0.3 and technique. Additional onsite hybridization and rapid 4.67 percent of all viable seed planted ultimately natural selection allow sustained establishment of emerge and develop into mature plants (Richardson adapted plant materials. and others 1986). Fourwing saltbush is widely planted in revegetation Fourwing saltbush is well adapted to interseeding. projects. When the seedings are carefully planted they Plants can be established in strips or furrows separate are usually successful. Evidence shows that fourwing from herbs (Giunta and others 1975; Ostyina and saltbush increases the value of otherwise herbaceous others 1984; Rumbaugh and others 1982; Stevens pastures (Ostyina and others 1984; Rumbaugh and and others 1981b). Interseeding is a useful means of others 1982; Ueckert and others 1990). Giunta and seeding the shrub into established stands of perennial others (1975) demonstrated the value of interplanting herbs. Narrow furrows or strips approximately 28 to fourwing saltbush in cheatgrass-dominated ranges. 30 inches (71.1 to 76.2 cm) in width can be created Sometimes, however, stands do not persist, perhaps using various interseeders (Giunta and others 1975; because of planting in sites that will not support the Stevens and others 1981b). Herbicides have also been species or because the source planted is not adapted to used for this purpose (Petersen and others 1990). the site. The species is a secondary or facultative Shrub seeds are placed in the clearing. Furrows can be selenium absorber and thus may be mildly poisonous interspaced at desired intervals, usually 10 to 12 ft where selenium occurs in the soil (Davis 1972; (3.0 to 3.7 m) apart. Fourwing saltbush grows pro- Kingsbury 1964). Minor losses have occurred when fusely and quickly produces a dense overstory. livestock had little or nothing else to eat (Hitchcock Fourwing saltbush grows well with understory herbs and others 1964). and other shrubs. When grown in association with In propagating fourwing saltbush, it is especially other species, herbage production of the fourwing important to be sure that the strain being planted is saltbush plants and the understory herbs remains adapted to the site. If possible, seed should be from a quite high (Rumbaugh and others 1981, 1982). source growing in a climate similar to that of the Fourwing saltbush is naturally susceptible to site being seeded unless other tested strains are basal decay disease in both plantation and natural known to have similar or better adaptation. Strains populations (Nelson and Welch 1984). This is not adapted to the mountain brush types, where soils usually a serious problem. Insects can be trouble- tend to be neutral, mature rapidly; but in deeper soils some to fourwing saltbush plants and populations, of this type, the life span has often been not more than especially when plants occur at heavy densities 10 years. On drier sites having higher alkaline con- (Haws and others 1984; Moore and Stevens 1984). In tent, life span of the same types has been much longer. seed orchards the case-bearing moth can be devastat- However, persistence of different accessions has been ing if not controlled. Malathion will effectively control quite erratic. Various biological and climatic influ- this pest (Moore and Stevens 1984). ences apparently affect persistence. Plants established from seed collected in the blackbrush-mesquite belt in

474 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 20 Chenopod Shrubs southern Utah have not persisted in colder parts of area. Other high polyploids (up to 20x), usually re- the State. Seed collected from colder areas have per- stricted endemics, occur in the southern portions of sisted well on warmer sites (Plummer and others the species range (Sanderson and Stutz 1994a; Stutz 1966, 1968). and Sanderson 1979). Tetraploids (4x = 2n = 36) are the common plants of the Intermountain areas. Varieties and Ecotypes Fourwing saltbush, with its outcrossing habit and common polyploidy, exhibits both allo- and autopoly- Fourwing saltbush probably has the greatest po- ploidy (Stutz 1978, 1984). Many of the common tential among the woody chenopods for reclaiming tetraploids of the Intermountain area appear to be arid ranges. It is one of the most widespread and autotetraploids (McArthur and others 1986). adaptable western shrubs. Fourwing saltbush is the Differences have been demonstrated among several center of a large species complex. It hybridizes widely accessions in winter protein content (Welch and Monsen with other woody saltbushes (Blauer and others 1976; 1981, 1984) and in both growth performance and Sanderson and Stutz 1994b; Stutz 1978, 1984). Sev- survival (Geist and Edgerton 1984; McArthur and eral subsidiary species have been recognized. These others 1983a; Shaw and others 1984; Young and oth- species are all fourwinged and likely have A. canescens ers 1984e). In planting fourwing saltbush, either a in their ancestry or are directly derived from that proven accession or accessions obtained from nearby species. Five of the allies of fourwing saltbush (A. or from higher latitudes or higher elevations than aptera, A. bonnevillensis, A. garrettii, A. navajohensis, the planting site should be used (McArthur and and A. “robusta”) are listed in table 3. Each species has others 1983a; Van Epps 1975). a habitat requirement different from A. canescens, and Three releases of fourwing saltbush of proven qual- consequently would be useful for restoration and re- ity and adaptability have been made (Carlson 1984). vegetation of particular kinds of sites (table 3). Additional work is under way to provide more plant Fourwing saltbush proper is composed of widely material with known characteristics by identifying occurring polyploid races (x = 9). Diploids are mostly and testing promising accessions and through hybrid- restricted to the southern portion of the species range— ization and selection procedures (Carlson 1984; Stutz New Mexico and Texas and south (Dunford 1984; and Carlson 1985). Stutz and Sanderson 1979). Hexaploids, too, are re- The three releases are ‘Rincon’, ‘Marana’, and stricted mostly to the west Texas and New Mexico ‘Wytana’. ‘Rincon’ is well adapted to a wide range of

Table 3—Other fourwing saltbushes in addition to Atriplex canescens.

Postulated Distinguishing Species origin Habitat Distribution characteristics References Atriplex aptera A. canescens Clay hill Upper Missouri Diminutive (or lacking) Brown 1956; x badlands River Drainage, utricle wings. Small Stutz 1978; A. gardneri Montana, Wyoming, 24 inches (60 cm) Stutz and the Dakotas plants others 1979. Atriplex bonnevillensis A. canescens Alkaline Eastern and Central Small 30 inches Hanson 1962; playas Great Basin, Utah (75 cm) highly Frischknecht and and Nevada variable plants Ferguson 1979. Atriplex garretti — Rocky, sandy Colorado River and Early flowering, Brown 1956; soils; in tributary canyons, succulent, yellow- Hanson 1962; canyons southern Utah, green leaves. Small Blauer and others northern Arizona 16 inches (40 cm) 1976. Atriplex navajoensis — Rocky, sandy Vicinity of Navajo Woody, yellow- Hanson 1962. soils Bridge, Coconino green leaves. County, AZ Up to 40 inches (100 cm) tall Atriplex “robusta”a A. canescens Highway East of Wendover, Rapid growth rate, Stutz and others x disturbances Tooele County, UT utricles rarely 1979. A. tridentata in salt flats winged but with variously shaped protuberances

aThis taxa has not been formally named.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 475 Chapter 20 Chenopod Shrubs soil textures: sandy areas, gravely washes, loamy Large, thick utricles must be mechanically frac- soils, heavy clay soils, and moderately saline soils. tured to attain uniform germination (Monsen and This strain is best adapted to big sagebrush and McArthur 1985). pinyon-juniper zones, but it also does well in the more mesic portions of the salt desert shrub area. It appears to have tolerance to salt similar to that of other Atriplex confertifolia ______fourwing saltbush ecotypes. Because of its high eleva- Shadscale tion origin, ‘Rincon’ has shown an adaptability to more northern climates than might be expected, consider- Shadscale is a compact spinescent shrub, growing its origin in northern New Mexico. It has pering typically in dense clumps from 0.5 to 4 ft high formed well at 3,000 to 8,000 ft (910 to 2,400 m) (15.2 to 121.9 cm) and 1 to 5.5 ft (30.5 to 167.6 cm) elevation where the average annual precipitation wide (fig. 5). The rigid, brittle branches are scruffy ranges from 9 to 23 inches (22.9 to 58.4 cm) (McArthur when young but become smooth and spiny with age. and others 1984b). ‘Rincon’ is being grown in seed The leaves are nearly circular to elliptic, oval, or orchards for commercial seed production (fig. 2b). The oblong, 9 to 25 mm long, 4 to 20 mm wide. They are recommended design and early orchard performance scruffy and gray especially at maturity (Blauer and characteristics of ‘Rincon’ are given by Noller and others 1976). others (1984). Flowers of shadscale are similar to those of four- ‘Marana’ is recommended for Mediterranean type wing saltbush except in the nature of the bracts climates below 4,000 ft (1,200 m) in elevations with enclosing the seed. The bracts of shadscale are foliose, minimum temperatures of 10 ∞F (–10 ∞C) generally in 5 to 12 mm long, broadly oval to almost round, united California and adjacent areas (Carlson 1984; Hassell at the base, and have entire, free, somewhat spreading 1982; McArthur and others 1984b). ‘Wytana’ is closely margins (fig. 5). Shadscale blooms from late March in related to A. aptera (table 3), and is adapted to eastern the southern portion of its ranges to mid-June in the Montana and Wyoming (Carlson 1984; McArthur and northern portion (Hanson 1962). Utricles mature about others 1984b). 15 weeks after blooming. They are fairly persistent Several other accessions may have utility for wide- through the winter months (Blauer and others 1976). spread planting although they haven’t been formally Although sex expression in shadscale has not been released (McArthur and others 1983a; Petersen and studied in the detail accorded fourwing saltbush, it others 1987; Shaw and others 1984; Young and others shares some of the characteristics of that species. For 1984e). One particular accession with a proven per- example, shadscale populations may exhibit biased formance record is the gigas diploid (up to 10 ft [3 m] sex ratios and include monoecious as well as pistillate tall) that grows naturally only in Utah’s Little Sahara and staminate individuals, and individual plants of Sand Dunes (Stutz and others 1975). This giant form different genders behave different physiologically layers readily and appears to have an important place (Freeman and Harper 1980; Freeman and McArthur in the stabilization of dunes or other sandy areas. In 1982, 1984). trials since the 1980’s, the giant form has grown well on clay, as well as sandy soils. Several other Utah accessions perform well in pinyon-juniper and big sagebrush communities and on mine disturbances throughout the Intermountain region (McArthur and others 1983a). Some selections or ecotypes are nearly evergreen; that is, they retain a high percentage of their leaves on a year- round basis. Ecotypes that have this feature occur in both the warmer and cooler portions of the plant’s natural range. Consequently, the feature is not restricted to limited areas. Among ecotypes considerable differences exist in seed size, utricle thickness, and germination patterns (Monsen and McArthur 1985; Springfield 1970b). Similar variation also occurs among seeds collected from a single bush. However, germination traits are distinct with different ecotypes, and the features can be used to increase stand establishment. Small utricles normally germinate more rapidly and uniformly than large utricles. Figure 5—Shadscale growing near Elberta, Utah County, UT.

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Ecological Relationships and Distribution Garvin and others (1996) using eight shadscale accessions concluded that the species germinates in a long- Shadscale is widely distributed in the Western term temporally staggered fashion, suggesting the United States from Chihuahua to northern Montana operation of multiple dormancy mechanisms. A few and from North Dakota and Texas to California and populations lack the species’ characteristic seed dor- Oregon (Branson and others 1967; Hall and Clements mancy. Insects can cause severe damage to shadscale 1923; Sanderson and others 1990; Stutz and Sanderson (Haws and others 1984; Hutchings 1952). 1983). In broadness of distribution among the woody Low seed fill often compounds low seed germination chenopods it ranks next to fourwing saltbush and problems. Seed is usually acquired from wildland winterfat and frequently grows in association with stands growing under arid conditions. Seed quality both. It grows at elevations from 1,500 to 7,000 ft varies from year to year and even among bushes. Most (460 to 2,100 m) usually on fine textured alkaline soils commercial lots of seed are collected from upland but also on coarser, sandier soils (Blaisdell and foothill ranges that receive higher amounts of pre- Holmgren 1984; Branson 1966; McArthur and others cipitation. Harvested seed crops are produced about 1978b). every 3 to 5 years, yet some sites are more reliable It occurs in nearly pure stands and in mixtures with than others. winterfat, budsage, big sagebrush, black sagebrush, Seed or utricles are normally hand collected but rabbitbrush, spiny hopsage, black greasewood, gray can be efficiently harvested with vacuum, rill, or strip- molly, horsebrush, juniper, other saltbushes, and sev- type machinery (see discussion under fourwing salt- eral species of grass. It endures alkaline soils better bush). Seed usually ripens in late fall or early winter. than most of these associates and can be found grow- Utricles must be dry before they detach from the bush. ing with such halophytes as glasswort (Hall and Early winter storms can interfere with seed har- Clements 1923). West, Caldwell, and their colleagues, vesting. Once harvested, the wings are usually re- (Caldwell and others 1977; Moore and others 1972; moved from the utricle with a hammermill or flail-type West 1985; West and Gasto 1978) showed how the beater. Seeds can be stored for a number of years synecological and physiological relationships of without loss of viability. However, unless stored in shadscale and winterfat growing in a mixed stand give cold chambers and treated with an insecticide, seed insight on why subtle environmental differences are borne insects can destroy stored lots. Good quality important in the distribution patterns of these and seed has between 40 to 50 percent “filled” utricles. other desert shrubs. A number of ecotypes grow on a However, a considerable amount of lower quality seed wide range of sites including highly alkaline soils. is sold. Seed viability should be retested if stored over Shadscale has a relatively short life span and can be 1 year. killed by drought and high water (Blaisdell and Because seed germination is often erratic and Holmgren 1984; McArthur and others 1978b). varies among ecotypes, fall seeding is recommended to aid in overcoming natural dormancy and break- Plant Culture down of the utricle. Seed must be placed 0.25 to 0.5 inch (6.4 to 12.7 mm) deep to ensure germination and Although shadscale readily spreads naturally from seedling establishment. Shadscale is often planted on seed, it is difficult to establish from seed. Only under heavy-textured salty soils. Soil particles may be floc- the most favorable climatic conditions do stands culated, which creates a rough surface that aids in become established from artificial seeding (McArthur seed burial. However, soils may crust easily, and and others 1978b; Monsen and Richardson 1984). It shadscale seedlings may not be able to emerge can be successfully transplanted when moisture through the surface crust. Shadscale seedlings grow conditions are right in the spring or fall (Luke and slowly and are not able to compete with herbs, par- Monsen 1984). Fruit remains on the ground 1 year or ticularly cheatgrass. Consequently, planting sites more before germinating and establishing. Under- must be cleared of weeds, and the shrub should be standing the riddles in shadscale germination and seeded alone in separate rows from other species. stand establishment requires more research. Warren Once seedlings reach 2 years old, losses to competi- and Kay (1984) demonstrated that dormancy in tion are low. The plant has done well on barren mine shadscale is difficult to overcome by traditional me- wastes, particularly infertile soils (Ferguson and chanical and chemical methods. Young and others Frischknecht 1985). (1984e), following a Stutz (1978) and Stutz and Broadcast seeding with no followup measure to Sanderson (1983), scenario of shadscale’s evolution in cover the seed has not been very successful. Either a pluvial lake regime, suggested that a great deal of direct drilling or interseeding is recommended. Seed ecotypic variation in shadscale germination should should be planted at approximately 2 to 4 lb PLS/acre occur. Oaks, Stutz, and Sanderson (unpublished) have (2.2 to 4.5 kg PLS/ha). Most conventional seeders can found ecotypic germination variation to be common.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 477 Chapter 20 Chenopod Shrubs be used to seed this shrub. Dewinged seeds are rela- readily seek this accumulation. In some areas shadscale tively small and flow freely through most seeding is an increaser and becomes dominant as more palat- devices. Seeds are relatively heavy and can be used in able species such as budsage and black sagebrush are seed mixtures to facilitate the planting of more fluffy killed by grazing (Blaisdell and Holmgren 1984). Be- or trashy seeds through the seeder (Monsen, unpub- cause shadscale is generally more salt-tolerant than lished). Seeding rates can be easily regulated. Seed fourwing saltbush and several other shrubby chenop- germination may begin quite early in the spring and ods, it may be used to better advantage on the more continue until early summer. salty areas for range rehabilitation and soil stabiliza- Natural spread of shadscale occurs erratically, al- tion (McArthur and others 1978b). Existing stands of though large stands frequently develop. New seed- shadscale should be managed rather than mechani- lings normally require 3 to 8 years to attain a mature cally manipulated because the species usually occurs status. Some ecotypes, however, from north-central on areas that are difficult to manipulate (Bleak and Nevada and northeastern Wyoming grow fast. Un- others 1965; Plummer 1966). This is because shad- doubtedly other collections have similar traits. scale normally grows in semiarid conditions, often occu- Transplants of shadscale can be successfully reared pying sites that receive less than 10 inches (25.4 cm) as container or bare root stock. Even under favorable annual moisture. When disturbed these areas are conditions, shadscale seedings grow rather slowly. usually invaded by cheatgrass or other weeds. The The 1 year old stock usually are 3 to 7 inches (7.6 to competition must be controlled to allow survival of 17.8 cm) in height but have a moderately diverse naturally or artificially seeded shadscale. Large ex- root system. tensive areas can be direct seeded if no competition Seedlings do not survive field planting well unless exists. Otherwise, some means of site preparation is extra care is given in handling and planting. Seedlings required to reduce competition. must be dormant when planted. Because some leaves Shadscale often occurs as near pure stands void of tend to remain on the plant, it is difficult to deter- most other shrubs. Plants from these pure stands do mine if plants are clearly dormant. Seedlings also not grow in mixtures with many other shrubs. Conse- require a growth medium that is similar to the origi- quently, they are usually planted alone. Ecotypes that nal soil. Bare root seedlings are particularly sensitive extend into pinyon-juniper woodlands and big sage- to overwatering. Maintaining plants as 2-0 stock re- brush communities have been more successfully seeded sult in much larger materials, but has not improved in diverse habitats. These ecotypes, the diploid races field survival. Transplants do not survive if planted (see Varieties and Ecotypes section), when seeded amid herbaceous competition. Clearings of 30 inches compete favorably with other species but are best (76.2 cm) in diameter are required to reduce competi- adapted to sites similar to those from where they have tion sufficiently. Field survival of 60 to 80 percent is been collected. Ecotypes collected from more arid salt considered normal for large-scale plantings. Much desert conditions, the higher polyploids, are less lower survival results if plants are planted offsite. adapted to foothill upland communities but appear Shadscale does not establish or persist long when more drought tolerant. planted on unadapted sites. Neither transplants nor direct-seeded seedlings persist if planted offsite. Care Varieties and Ecotypes should be taken to assure seeding of adapted ecotypes. Unlike fourwing saltbush, shadscale seedlings and Shadscale comprise chromosome races ranging young plants are not selectively sought and grazed by from diploid (2n = 18) to decaploid (2n = 90). The rodents, rabbits, or livestock. chromosome races are distributed, in general, with the high ploidy levels occurring in uniform stands on Uses and Management valley floors and lower ploidy forms progressively larger and more morphologically variable and occur- Despite its spiny character, shadscale is grazed by ring in more diverse communities (Sanderson and livestock and game. Persistence of leaves varies others 1990; Stutz and Sanderson 1983). Little atten- greatly over shadscale habitat. Forage value improves tion has been paid to differential adaptation of these with persistence of leaves. Spines save this species forms for revegetation purposes. However, the most from heavier grazing than it would otherwise receive, robust erect forms produce more seed and have gener- and the spines are likely a factor in its common ally been used more for seeding. More attention should widespread distribution. Seeds are a preferred and be paid to this feature because some of the higher nutritious part of the plant (USDA Forest Service polyploids are narrow genetically and adaptively. Con- 1937). According to Sampson and Jespersen (1963), as sequently, these are poorly adapted to many sites for the leaves and fruits drop in the autumn, they often which the more diverse diploids and tetraploids are accumulate around the parent shrubs. Livestock adapted (Stutz 1982; Stutz and Sanderson 1983).

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Atriplex corrugata ______of salt occur (Hanson 1962). Mat saltbush is probably the most halophytic shrub in the genus, but it is Mat Saltbush frequently found in areas where the salt concentration is not so high. In these areas, it may be associated with Mat saltbush is a low shrub that forms dense, shrubs such as winterfat, cuneate saltbush, shadscale, extremely prostrate (6 inches [15.2 cm] tall) nearly fourwing saltbush, black greasewood, budsage, and white mats five to 20 times wider than they are tall gray molly. Young plants have been successfully trans- (fig. 6). The prostrate branches often produce adven- planted into a former greasewood type in the Great titious roots where they contact the soil. The bark is Basin (Blauer and others 1976). soft, spongy, and white. The evergreen sessile leaves are opposite on the lower parts of the stems, and Plant Culture alternate above. The blades are densely scurfy and measure 7 to 18 mm wide (Blauer and others 1976; The seed or utricle does not require hammermilling. McArthur and others 1978b). Bushes are commonly The seed is usually screened with a fanning mill to dioecious, but some are monoecious. The sexual sys- remove debris, and then seeded with most conven- tem appears to operate much like that of fourwing tional equipment. Cleaned seed can be stored for and shadscale (Freeman and McArthur 1984; Pope several years without serious loss in viability. Most 1976). The yellow to light-brown staminate flowers native stands produce between 40 to 60 percent filled are borne of glomerules 3 to 6 mm wide on nearly utricles. naked spikes. The pairs of fruiting bracts that en- Seed production is often quite low, yet even during close the pistils of the female flowers are sessile or serious drought and prolonged use, some shrubs pro- subsessile, 3 to 5 mm long, 4 to 6 mm wide, united duce seed. Seed fill or quality is usually low. Some along seven-tenths of their length and usually densely stands, particularly those in south-central Wyoming, tuberculate on the lower one-third. Mat saltbush produce unusually abundant crops yearly. Large flowers from April to June, and the fruit ripens 6 to fields are machine harvested and sold each year. 10 weeks later. Hand collection is quite slow, yet field combines are able to harvest seed easily. Ecological Relationships and Distribution Uses and Management Mat saltbush is distributed mainly on soils derived from the Mancos shale formation in eastern The plant is important as a winter forage. It often Utah, southwestern Wyoming, western Colorado, grows interspaced with other plant communities at and northwestern New Mexico at elevations from mid to high elevations. Under these circumstances it 4,000 to 7,000 ft (1,200 to 2,100 m). It tolerates up to is encountered on exposed shaley outcrops with few 13,000 ppm soluble salts and is often the only peren- other plants. These sites are often windblown slopes nial plant present where such high concentrations that are accessible to winter grazing. The shrub withstands heavy use, even under unstable soil conditions. Plants may be heavily browsed and seriously trampled. Big game animals often concentrate on these exposed sites during periods of deep snow accumulation. The shrubs are able to recover from serious abuse without noticeable loss of vigor. Fall seeding is normally recommended. Where possible, midwinter planting is successful. Seedlings are susceptible to early spring frosts. If serious frosts are expected, spring seeding can be used. Seeds should be planted near the soil surface or not more than 0.5 inch (1.3 cm) deep. Seeding is usually done on sites that once supported the shrub. These areas usually are not infested with annual weeds, consequently, direct seeding without other preparatory measures is all that is necessary. Seedlings usually emerge rapidly and grow rather vigorously the first year. If moisture remains avail- Figure 6—Male mat saltbush (left) and able, seedlings normally continue growth all summer. female Castle Valley clover saltbush (right) Mat saltbush has been seeded with selected grasses growing along highway near Fremont including Russian wildrye, tall wheatgrass, and crested Junction, Emery County, UT. wheatgrass. Although these herbs may improve the

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 479 Chapter 20 Chenopod Shrubs forage production, they usually do not persist in dense elliptic evergreen leaves are 2 to 6 cm long and 0.5 to stands throughout the site. Young grass seedlings 2.5 cm wide (Hall and Clements 1923). In common and mature plants do suppress shrub establishment. with some other woody saltbush species, this species is However, grasses interseeded into established shrub apparently trioecious (Freeman and McArthur 1984). stands do not decrease shrub density or vigor. Its yellow to brown staminate flowers are borne in Mat saltbush can be broadcast seeded and covered glomerules arranged in panicles. The pistillate flow- with a drag or harrow if planted alone. When seeded ers are borne in axillary clusters and consist of pistils with herbs, the shrub seeds should be planted in enclosed by wingless bracts. At maturity, the bracts separate rows or spots. Between 2 to 4 lb PLS/acre are 5 to 9 mm wide, irregularly toothed along their (2.2 to 4.5 kg PLS/ha) is recommended. However, margins, and have numerous, flattened, crestlike tu- satisfactory stands have developed from lower rates. bercles on their sides (fig. 4d). Blooming occurs from When seeding on the heavy clay soils that normally mid-April to July, depending on elevation and climatic support this shrub, care must be taken to avoid surface conditions. Ripening of the fruit follows about 7 weeks compaction and crusting. later (Hanson 1962). Transplanting either nursery-grown bare root stock, wildings, or container materials is successful. Wild- ings can be dug from native stands and transplanted Ecological Relationships and Distribution directly on disturbances. Wildings and bare root stock The species occurs in high to moderately alkaline survive abusive treatment, but should not be unneces- soils in eastern Utah, southwestern Colorado, and sarily mistreated. Transplants should not be allowed northern New Mexico. In these areas, Castle Valley to “heat up,” and roots must be placed in a moist soil. clover saltbush is often dominant or is codominant When transplanted on unstable and seemly infertile with shadscale and mat saltbush. soils, the transplants survive remarkably well. Natu- ral enlargement of the crown occurs soon after planting, providing excellent soil protection even under Plant Culture adverse conditions. Seed production of Castle Valley clover saltbush is Mat saltbush has performed well from either direct similar to that of other woody saltbushes. Annual seeding or transplanting disturbed sites. It is par- yields are erratic, and only about 50 percent of all ticularly useful as a shrub for seeding mine distur- utricles produce viable seeds. Seed germination is bances, road construction sites, and similar areas also erratic and varies among ecotypes. Seeds or where heavy textured soils and substrata are exposed. It has established, persisted, and grown well when utricles are relatively small and can be planted with planted on pinyon-juniper and big sagebrush sites if most equipment. Seedlings are quite vigorous and soils are quite basic and heavy. grow moderately rapidly. Young plants are competi- Small seedlings normally are not destroyed by graz- tive but can be suppressed by perennial grasses and ing or insect attack. The shrub should be seeded in annual weeds. Plants invade open disturbances, yet areas where remnant plants remain. Attempting to success from artificial seeding is not consistent. The convert mat saltbush communities to fourwing salt- plant should not be seeded directly with herbaceous bush, winterfat, or blackbrush is not advised. Al- plants but is able to compete favorably when seeded though these and other shrubs may be more palat- with fourwing saltbush, big sagebrush, green ephe- able, converted communities usually do not remain. dra, and spiny hopsage. The shrub should not be seeded on areas other Varieties and Ecotypes than sites naturally occupied by this shrub. The species is not well adapted to the big sagebrush and Little study has been accomplished in this area. upland communities, but can be used throughout Hanson (1962) and Pope (1976) have demonstrated most salt desert shrublands. Like other woody salt- some populational differences. Mat saltbush will hy- bushes, the shrub does well on fresh disturbances and bridize with other saltbush species (Blauer and others can be used to treat exposed substrata created from 1976; Stutz 1978, 1984). mining, road construction, and so forth.

Atriplex cuneata ______Uses and Management Castle Valley Clover or Cuneate Saltbush Castle Valley clover saltbush, so named by stock- men because of the high livestock preference in all Castle Valley clover saltbush is a low shrub 0.3 to seasons, remains green and succulent throughout 1.5 ft tall (9.1 to 45.7 cm) with a more or less prostrate, the winter. The shrub shows particular promise in woody, much-branched base and erect branches artificial restoration of certain winter game ranges (fig. 6). The light grey-green, spatulate to broadly and on disturbed sites having saline-alkaline soils.

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It grows through a wide range of salinity and may be (Hanson 1962; Pope 1976). The brown staminate associated with any of the other shrubby chenopods flowers are borne in glomerules 3 to 5 mm wide on and many of the sagebrush species. It grows well with nearly naked one-branched terminal panicles. The a variety of desert grasses and forbs. Its protein pistillate flowers are borne on leafy spikes. Fruiting content helps to provide a balanced diet for livestock bracts that enclose the female flowers are 3 to 6 mm and game, particularly in the winter when much of long, 2.5 to 5.0 mm wide, and are sessile. Their the associated vegetation is dry (McArthur and others surfaces range from smooth and free of tubercles to 1978b). densely tuberculate. The apex of the bract ends in an oval terminal tooth subtended by two slightly smaller Varieties and Ecotypes lateral teeth. Gardner saltbush flowers from mid- May to the first of July and intermittently following Castle Valley clover saltbush hybridizes readily heavy rains. The fruit ripens about 7 weeks after with a number of the saltbushes, but particularly so flowering. with fourwing saltbush, both naturally and artificially (Blauer and others 1976; Stutz 1984). A stabilized Ecological Relationships and Distribution hybrid of these could be a boon to our winter ranges. Such a plant would be taller than Castle Valley clover Gardner saltbush is abundant throughout much of saltbush and retain more green foliage than four- Wyoming and Montana on clay and occasionally sandy wing. A stabilized, widely adapted hybrid might be soils. It is tetraploid, 2n = 36 (Stutz and others 1979). found in nature. It is useful year-around forage for big game and livestock (McArthur and others 1978b).

Atriplex gardneri ______Plant Culture Gardner Saltbush Field establishment may be difficult because of low seedling vigor (Ansley and Abernethy 1984a,b). Ansley Gardner saltbush is a low subshrub, usually less and Abernethy (1984b) demonstrated that plants are than 12 inches (30.5 cm) (fig. 7). The lower one-fourth best established in a field situation when seeds are to three-fourths of the plant is slightly woody and the scarified and stratified. rest is herbaceous. Its habit varies from decumbent For production fields and commercial harvest, creeping forms to rounded forms three to five times Carlson and others (1984) recommend direct seeding broader than they are high. The spineless decumbent on a firm weed-free seedbed with 36 inch (91.4 cm) branches usually produce adventitious roots where rows in the fall at a rate of 20 pure live seed per foot they contact the soil. The lightly scurfy leaves are (66 per m). Once the plants are established and pro- evergreen spatulate to oblaceolate to obovate 15 to ducing seed, the plants can be cut by hayswather, 55 mm long, and 5 to 12 mm wide. Plants of this windrowed, and the utricles collected by conventional species are dioecious or sometimes monoecious grain combine.

Uses and Management Gardner saltbush can be successfully seeded on mine disturbances and rangelands. Although initial establishment is often erratic, a slow but general improvement normally results. When seeded in areas free of weedy competition, seedling survival is usually adequate. Shrub seedlings grow quite favorably and are able to survive normal grazing pressure.

Gardner Saltbush Relatives ______Atriplex buxifolia, Atriplex falcata, Atriplex tridentata Much of the literature dealing with Gardner saltbush is confusing in that there are several related Figure 7—Gardner saltbush (right) growing with species considered as a species complex. The taxo- budsage (left) in the Red Desert, Sweetwater nomic complexity has not been completely unraveled. County, WY. The species complex has been alternatively treated

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 481 Chapter 20 Chenopod Shrubs as the A. gardneri or the A. nuttallii complex. These suffrutescent shrubs exhibit population variability, hybridization between populations of the same and different taxa, and polyploid chromosome races (Stutz 1978; Sanderson and Stutz 1994b). Atriplex nuttallii is a name that has recently been lectotypified from the northern Great Plains (Saskatchewan) by Bassett and others (1983); that name (A. nuttallii) had been without standing prior to then (Fosberg 1941; Stutz and others 1979). We believe the plants of the Intermoun- tain area including A. buxifolia, A. falcata (falcate saltbush), and A. tridentata (tridentate saltbush) are best referred to as part of the A. gardneri complex reserving A. nuttallii for plants of the northern Great Plains. Drs. Howard Stutz and Stewart Sanderson (n.d.) are actively working on this group of plants from Figure 8—Example of root sprouting in trident a cytogenetic and morphological standpoint. They saltbush from west Sanpete Valley near Ephraim, propose the name of A. canadensis as an alternative Sanpete County, UT. name for A. nuttallii. Atriplex buxifolia is similar to A. gardneri in habitat preference (heavy clay soils) and in morphology (Hanson 1962; McArthur and others 1978b). Stutz Research with A. tridentata and A. falcata have (personal communication) believes that there is no been hampered by poor seed production. Although good reason to maintain the two as separate species both species are capable of good seed yields, concen- but to refer both to A. gardneri. Plants that have been trated efforts have not been developed to acquire referred to as A. buxifolia were larger than A. gardneri adequate samples. Both species have been success- (up to 50 cm; 19.7 inches) and distributed more to the fully established by fall seeding. The shrubs appear north and east (northern Nebraska to southern capable of growing over a wide range of sites, includ- Saskatchewan) (Hanson 1962; McArthur and others ing areas naturally occupied by fourwing saltbush. 1978b). The erect growth habit of A. tridentata and A. Both A. tridentata and A. falcata are generally falcata can be grown as bare root or container trans- larger statured than A. gardneri. Atriplex tridentata plant stock. They survive field planting quite well. plants may be from 4 inches to 3 ft tall (10.2 to 91.4 cm). Individual plants grow well under nursery conditions They root sprout profusely (fig. 8). This is the plant and may require pruning to reduce plant size. referred to as A. gardneri by Nord and others (1969) Atriplex tridentata and A. falcata often lack a herba- (see Stutz and others 1979). Atriplex tridentata can ceous understory. This is often due to heavy grazing. tolerate salt concentrations of over 30,000 ppm and Both shrubs persist under intense use, although grows with iodine bush and inland saltgrass seed production is reduced. Both shrubs are capable (Hanson 1962). It is distributed mostly in the Lake of growing with a diverse understory. These shrubs Bonneville Basin of western Utah, eastern Nevada, have the unique ability to persist with various forage and southern Idaho, but also in eastern Utah and species, and yet remain quite productive themselves. southern Wyoming (Hanson 1962; Stutz and others 1979). Atriplex falcata grows up to 20 inches (50.8 cm) Other Saltbushes ______tall. It is distributed through the northern Great Basin extending northward to eastern Idaho and Atriplex lentiformis (quailbush), eastern Oregon. It grows in soils with 9 to 50 percent Atriplex obovata (broadscale saltbush), sand often between or in association with shadscale, Atriplex polycarpa (allscale saltbush), Wyoming big sagebrush, and winterfat communities Atriplex hymenelytra (desert holly saltbush) (Hanson 1962; Yensen and Smith 1984). Atriplex tridentata is mostly hexaploid, but tetraploids are also Some other saltbush species occur in parts of the known (Stutz 1978; Stutz and others 1979). All the Intermountain area—especially in the southern saltbushes treated in this section are useful year- parts. Quailbush (Atriplex lentiformis) is a large around forages as is A. gardneri itself. They may shrub up to 11.5 ft (3.5 m) tall and 24.5 ft wide (7.5 m) prove useful in rehabilitation of disturbed saline sites. (fig. 9). Its leaves are evergreen, alternate, gray-green, The edaphic adaption of each taxa should be borne in scurfy. The distinctive utricles are small flat discs mind in rehabilitation efforts. about 3 to 4 mm in diameter. The similar species,

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are elliptical to obovate, 1 to 3.5 cm long, 1 to 2 cm wide, with a short petiole. This species has yellow staminate flowers borne in small glomerules and arranged 500 to 5,000 to a panicle. The fruiting bracts are broadly cuneate or obovate, 4 to 5 mm long, 5 to 9 mm broad, with smooth or sometimes slightly tuberculed surfaces (Blauer and others 1976). Broadscale saltbush tolerates salinity from 165 to 4,900 ppm soluble salts (Hanson 1962). It is commonly associated with such halophytes as black greasewood, seepweed, and alkali sacaton, and other saltbushes (McArthur and others 1978b; Wagner and Aldon 1978). Broadscale saltbush is an important browse plant Figure 9—A large quailbush saltbush near in alkaline areas (Edgar and Springield 1977; Hall St. George, Washington County, UT. and Clements 1923). It grows rapidly and is most succulent in the spring. Within areas of adaptation, this species should be useful for stabilizing disturbed sites. Frischknecht and Ferguson (1984) reported Torrey saltbush differs by being smaller, up to 10 ft that broadscale saltbush was the most successful of (3.0 m) tall and 16 ft (4.9 m) wide, and in having anled seven chenopod shrubs transplanted into a processed stems (versus terete for quailbush). Both species oil shale matrix in eastern Utah. After 6 years, over occur in the St. George area of Utah and in other 80 percent of the shrubs survived in both irrigated southern and western margins of the Intermountain and nonirrigated plots. Broadscale saltbush seed area (Brown 1956; Hanson 1962). One selection, ‘Casa’, germinates best following an after-ripening period of quailbush has been released for upland game use (Edgar and Springfield 1977). It has been observed to and erosion control (Carlson 1984). In contrast to some invade the disturbed margins of newly constructed other Atriplex species, quailbush utricles require burial highways. for effective plant establishment (Young and others Allscale saltbush is an intricately branched, rounded 1980). shrub to 5 ft (1.5 m) tall. Its twigs are delicate, even Three other southern saltbush species are broad- approaching spininess. Leaves are obovate to linearly scale saltbush, allscale saltbush, and desert holly obovate. It occurs in the Sonoran and Mohave Deserts saltbush. Broadscale saltbush is a subshrub 0.6 to 1.5 and in drier portions of the San Juaquin and Owens ft (0.15 to 0.45 m) in height with a woody, spreading Valleys of California. This species has value as browse base that produces numerous ascending to erect and as upland game habitat (Brown 1956; Chatterton branches (fig. 10). The silvery, scurfy, deciduous leaves and others 1971; Hall and Clements 1923; Sankary and Barbour 1972). Chatterton and others (1971) reported that allscale saltbush is an excellent nutrient source for browsing animals especially late in the year. Graves and others (1975) reported that allscale saltbush was useful for revegetating Mohave Desert land. For successful reseedings, they suggested that allscale saltbush be drill seeded in the early winter. Sankary and Barbour (1972) reported that November is the time of natural seed dispersal and seedling establishment for allscale saltbush. Mixing activated carbon directly with utricles in the seedbox enhances germination by inactivating inhibitory substances found in the utricles (Graves and others 1975). Desert holly saltbush is a compactly branching, rounded shrub to 3 ft (91.4 cm) tall. The twigs are brittle but not spiny. Its leaves are obicular in outline, but leaf margins are irregularly undulate or coarsely toothed and as long as 1.6 inches (4 cm). Desert holly Figure 10—Broadscale saltbush growing in is distributed on dry alkaline alluvial fans and hills Hamblin Wash, Coconino County, AZ. Ruler is from Owens Valley, California, to southwestern Utah 12 inches (30 cm) long. and south to Mexico (Brown 1956).

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Ceratoides lanata ______Ecological Relationships and Distribution Winterfat Winterfat is remarkably resistant to drought, and even on dry sites will produce seed in the third and Winterfat is an erect or spreading subshrub that fourth years after establishment. On favorable sites, shows wide variation in stature from dwarf forms plants have produced some seed in their first year of less than 8 inches (20.3 cm) in height to larger forms growth. The species has a deep taproot and numerous to 4 ft (1.2 m) (fig. 11). The dwarf forms are herbaceous extensive lateral roots. The plant may appear dead above a woody base. Taller forms tend to be woody after unusually dry years, but it normally recovers throughout. Branches and leaves are covered with a after rain. It exhibits strong reproductive qualities; dense coating of stellate and simple hairs that are however, establishment from artificial seeding is hin- white when young but become rust colored with age. dered because seedlings are highly sensitive to frost The leaves are alternate, linear, 5 to 50 mm long, damage (Stevens and others 1977a). with entire, strongly revolute margins. Winterfat is Winterfat stands often attain old ages. For example, predominately monoecious, but dioecious plants also in southern Idaho a stand of winterfat had plants may occur. The flowers are borne in dense paniculate thought to be an average of 72 years old, whereas the clusters along the upper portion of the branches. The corresponding figure for falcate saltbush was 24 years pistillate flowers are below the staminate flowers on (Yensen and Smith 1984). Determining the ages of monoecious plants. Staminate flowers lack bracts chenopod shrubs is difficult and imprecise (Stewart and petals. They have four sepals and four stamens and others 1940; Yensen and Smith 1984). borne opposite the sepals. Pistillate flowers lack both Winterfat is most abundant on lower foothills, sepals and petals. Pistils are enclosed by a pair of plains, and valleys with dry subalkaline soils in Utah, bracts that are united more than half their length. Nevada, Wyoming, Arizona, and New Mexico (Riedl The bracts are covered and often obscured by long, and others 1964; Stevens and others 1977). In the silky hairs (fig. 4i). These long hairs distinguish win- Great Basin, it often occurs in pure stands over terfat from species of Atriplex. Site and climate per- thousands of acres (Benson and Darrow 1945) and is mitting, winterfat blooms between May and August. an important component over millions of acres of the The fruit ripens from September to November and is salt desert shrub type. It also grows intermixed with dispersed by wind in late fall and winter. Seed pro- basin big sagebrush, pinyon-juniper, and ponderosa duction is extremely variable. A scant crop is pro- pine. The ecotypes for these upland communities duced in most years under arid circumstances. have been used to seed areas often lacking this Sources from foothill ranges usually produce an ad- shrub. Winterfat ranges from Canada through the equate crop each year. A heavy seed crop can be Great Basin and Rocky Mountain States to Mexico, produced in years of good summer storms coupled from California and eastward to Texas and North with the absence of summer grazing. Such productive Dakota (Branson 1966; fig. 1). It also grows over a wide crops on dry desert ranges may occur but once in a range of altitudes. In Utah, it occurs from the lower decade (Blauer and others 1976). Sonoran zone to the alpine ridges. Dwarf forms usually occur on desert floors, on areas of high salt concentration, and on high mountain tops. The larger forms occur on alluvial fans, foothills, and mesas on ponderosa pine and juniper-pinyon sites.

Plant Culture Winterfat can be established in seed production fields. To do so requires a firm, weed-free seedbed. Carlson and others (1984) recommended direct seeding at a depth of 0.12 inch (3.2 mm) in 36 inch (91.4 cm) rows. Fields require cultivation or hoeing and may require irrigation. Seeds can be combine harvested, usually from late October to mid-November. See chapter 24 for additional information on seed harvest. Seed is often hammermilled to remove the fluffy utricles (Carlson and others 1984). Booth (1982), however, reported better establishment by seeding whole Figure 11—‘Hatch’ winterfat growing at Bell Rapids, fruits on the soil surface. Hammermilling also dam- Gooding County, ID. ages many fruits. However, a problem with not

484 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 20 Chenopod Shrubs hammermilling is that fruits may be difficult to Seedlings of few other shrubs survive and grow as well separate from stems, leaves, and litter collected in as winterfat amid competition. However, seedlings the field. Seed should be cleaned using a barley can be weakened by competition, and weedy annuals debearder to detach separate utricles from the flower and established perennials can and do reduce seeding head, and then processed with a fanning mill to sepa- survival. Winterfat can be established by broadcast rate utricles from debris. seeding directly with herbs and other shrubs. When Winterfat can be successfully seeded and trans- drill seeded it is best to plant winterfat in separate planted in rehabilitation plantings (Clary and furrows from the herbs to reduce competition and to Tiedemann 1984; Frischknecht and Ferguson 1984; set the furrower depth for proper seed placement. Luke and Monsen 1984; Plummer 1977; Shaw and Winterfat establishes well when seeded with a culti- Monsen 1984; Stevens and others 1981b). When seeded packer seeder. Although seeds need only shallow alone, winterfat seed does not flow through a drill or coverage, planting in a firm seedbed is recommended. other seeding equipment. If mixed with heavier seed Young plantings are extremely persistent and are able or a carrier, the winterfat seed is forced through the to withstand heavy grazing, trampling, and drought. machinery. Even when mixed with heavier seeds, Winterfat is unusually well-adapted to mixed winterfat debris can cause bridging and clogging. seedings, particularly when planted in pinyon-juniper Seed must be carefully cleaned to facilitate seeding. communities. Although desert types normally grow Sticks and other debris should be removed by screen- alone, the tall robust ecotypes do well when seeded in ing. Chopping or hammermilling leaves a consider- mixtures. However, to encourage spread, openings able amount of material that restricts seeding opera- must be created to allow seedling survival. tion. Pelleting the seed has improved seeding without Winterfat is easily grown as container or bare root reducing seed germination. The heavy coated seed can nursery stock. Seedlings are susceptible to damping also be broadcast seeded with less chance of drift. off organisms, and entire plantings can be eliminated However, coated or pelleted seed must be properly in a few days. Maintaining clean, disease-free condi- cleaned prior to treatment, or sticks and leaves can tions is the best control measure, although soil drenches still impede seeding. and fumigation can be helpful. Young seedlings trans- Winterfat seed maintains viability only for rela- plant well. Plants are difficult to “harden-off” prior to tively short periods (6 months to 4 years) without field planting. Transplants that are not hardened special treatment (Kay and others 1977d; Stevens and are susceptible to stress by drought and freezing. others 1977, 1981a). Special treatments—such as use Spring planting is usually recommended, yet trans- of desiccants, sealed containers, and cold storage— plants are extremely hardy and survive even midsum- can prolong viability up to 8 years (Kay and others mer planting. The stems of young seedlings become 1977d; Springfield 1974). Winterfat seed requires an quite brittle and can be easily broken off during lifting after-ripening period for maximum germination and and transplanting. germinates best at warm temperatures (77 to 80 ∞F [25.0 to 26.7 ∞C]) (Springfield 1968, 1972b; Young and Uses and Management others 1984a). Current research seeks ways to better establish winterfat by seeding because seedlings are Winterfat is a superior nutritious winter browse for generally vulnerable to spring frosts (Stevens and livestock and big game. Sheep, cattle, antelope, elk, others 1977a). deer, and rabbits consume it. Except for the woody base Winterfat seed is usually cleaned to a purity of and larger stems, the plant is edible (McArthur and between 20 to 50 percent. Seed viability normally others 1978b; Riedl and others 1964). Overgrazing has ranges between 20 to 50 percent. Approximately 1 lb greatly reduced and even eliminated winterfat in some (PLS) per acre (1.1 k/ha) is recommended for drill areas, even though it is relatively tolerant to grazing seeding. Seeds can be successfully established by broad- (Stevens and others 1977a). Rasmussen and cast seeding, if lightly covered with a drag, harrow, or Brotherson (1986b) demonstrated that late winter pipe harrow. Seeds should be covered with not more grazing of winterfat by cattle reduces the vigor of than 0.25 inch (6.4 mm) of soil. Some seedlings estab- winterfat stands and opens those stands to less lish from surface seeding, but failure to cover the seed desirable forage species. Winterfat is potentially one is a major reason for poor stands on semiarid sites. of the most useful shrubs for planting to increase cover Because spring frosts seriously weaken young seed- and forage on alkaline soils of desert ranges in Utah and lings, spring plantings are recommended in areas adjacent States where the average annual precipita- where spring frosts are expected. tion is less than 10 inches (25.4 cm). It is one of a few Winterfat germinates quickly and seedlings grow shrubs that can be reliably seeded under arid circumstances. A great deal of winterfat is planted on West- rapidly. These traits, coupled with early germination, ern ranges. Plummer (1984) estimated that 20,000 lb cause the plant to be highly competitive with herbs.

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(9,100 kg) of seed were sold annually. Winterfat ecotypes Grayia brandegei ______from upland communities have proven useful for improving deteriorated pinyon-juniper game winter Spineless Hopsage ranges. Sampson and Jespersen (1963) give winterfat a Grayia, like Atriplex, has staminate flowers of four browse rating of excellent to good for cattle and or five sepals, four or five stamens, and pistillate sheep, excellent to fair for goats, good to fair for flowers that lack both sepals and petals but have deer, and fair for horses. Shaw and others (1984) pistils enclosed by pairs of bracts. The margins of showed that some winterfat ecotypes have potential as each pair of bracts are united from base to apex except upland game cover. Ostyina and others (1984) re- for a minute apical opening. Pendleton and others ported that winterfat is useful in increasing range (1988) have shown that spineless hopsage has the productivity for fall and winter use of grass pastures. unusual monoecious floral system of heterodoch- The species is a good natural increaser and should be ogamy. In that system protandrous (“male” first) and highly useful for stabilization in areas where it natu- protogynous (“female” first) plants are synchronized rally occurs. Some good stands have resulted from and reciprocal ensuring cross fertilization. On the broadcasting utricles on such areas (McArthur and basis of a morphological, embryological, and chro- others 1978b). matographic study, Collotzi (1966) recommended that Grayia be reclassified as Atriplex. Welsh (1984) made Varieties and Ecotypes an alternate suggestion that G. brandegei be transferred to Zuckia as Z. brandegei var. brandegei. We The different ecotypes of winterfat not only show favor retaining Grayia as a genus including both the wide variation in stature but also in seed production, species G. brandegei and G. spinosa. McArthur (un- seed size, seedling germination and vigor, pubescence published) analyzed Collotzi’s (1966) chromatographic on fruit and seed, and tolerance to varying pH in data, which shows G. brandegei and G. spinosa to be soils. These ecotypes maintain character differences most similar among the 10 chenopod taxa. Zuckia was when planted in uniform gardens (Riedl and others not included in Collotzi’s (1966) study. More work is 1964; Stevens and others 1977a). A more woody, some- required to resolve the Grayia-Atriplex-Zuckia taxo- what spinescent form of winterfat (C. lanata ssp. nomic question. subspinosa) referred to as foothills winterfat occurs on Spineless hopsage is a subshrub to 3.5 ft (1.1 m) tall rocky hills in southern Utah, Arizona, California, and (fig. 12) with linear-oblanceolate to obovate leaf blades Mexico. It apparently is the only form present in 1.5 to 4.5 cm long (Collotzi 1966; fig. 13). The flowering southern Arizona. This subspecies shows intergrada- period varies between mid-June and mid-August. tion in all characteristics with typical winterfat Seed matures in late September through early October. (Kearney and Peebles 1960). Six other species of Utricles persist on the plants until January and some Ceratoides are confined to the old world (McArthur may last through the winter. These are often removed and Sanderson 1984). by small mammals and birds. Some controversy still exists as to whether the long standing name of Eurotia retains its validity in Eur- Ecological Relationships and Distribution asia and in North America (Committee for Spermatophyta 1978; Howell 1971; Reveal and Spineless hopsage is generally restricted to shale formations of the Upper Colorado River Drainage Holmgren 1972; Robertson 1982). There is also some (Stutz and others 1987) in southwestern Wyoming, discussion that the proper generic name is western Colorado, northwestern New Mexico, north- Krascheninnikova. Robertson (1982) suggested that eastern Arizona, and eastern Utah. Other popula- south-central Asian winterfat, also known as Pamirian tions occur in central Utah. It favors silty clay loam winterfat (Ceratoides latens or Eurotia ceratoides) has soil derived from shales. It grows on tough sites that value as a forage plant on North American ranges. few other plants occupy (Pendleton 1986; Pendleton ‘Hatch’ winterfat is a recently released selection and others 1988). Collotzi (1966) noted it occurred that is suitable for its ability to establish, persist, and under mildly alkaline (pH 7.4 to 7.7) conditions. provide forage in sagebrush and pinyon-juniper communities (Monsen and others 1985a; Stevens and Plant Culture Monsen 1988). ‘Hatch’ is a large ecotype capable of providing winter forage available above the snow Seed viability and production is similar to saltbush (fig. 11). This selection also has potential for upland species. Low viability is often encountered due to the game habitat (Shaw and others 1984). arid conditions at the native sites. It germinates and produces seedlings readily, but unless protected, these are quickly taken by rodents and rabbits.

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Figure 12—Spineless hopsage growing in Figure 13—Spineless hopsage growing in Antelope Valley, Sanpete County, UT. Antelope Valley, Sanpete County, UT.

Uses and Management are similar. It has small, inconspicuous, unisexual flowers. The sexual system appears to be temporally This species may be used as browse by livestock monoecious to allow the species to more efficiently and big game. Small mammals have shown a high occupy stressful, heterogeneous, patchy environments preference for seedlings established at several loca- (McArthur and Sanderson 1984). Spiny hopsage is an tions. The shrub should be particularly useful for erect, diffusely branched, spinescent shrub from 1 ft stabilizing disturbed shaley soils. It provides forage (30.5 cm) to 4 ft (121.9 cm) high. Its leaves are fleshy for livestock and game, especially in the spring in the spring but usually dry before midsummer. when it is full leaf (McArthur and others 1978c). Fruits (fig. 4k) generally mature just before leaf fall Kingsbury (1964) listed spineless hopsage as possible secondary or facultative selenium absorber. As such, (Ackerman and others 1980). Unusual ecotypes in it could be mildly poisonous in areas where the soil northeastern Utah and southwestern Idaho retain a contains selenium. Some ecotypes demonstrate the considerable percentage of their leaves throughout ability to grow on heavy soils within pinyon-juniper the growing season (Blauer and others 1976). and basin big sagebrush communities. Stands establish best from fall plantings. To date the shrub has Ecological Relationships and Distribution not grown well with other herbs or shrubs. However, it is well suited to harsh situations. Seedlings grow Spiny hopsage occupies plains, desert valleys, and quickly and can be reared as transplant stock quite foothills over much of the Western United States easily. (eastern Washington south to southern California and east to western Wyoming and northwestern Varieties and Ecotypes New Mexico) at elevations ranging from 2,500 to 6,900 ft (760 to 2,100 m) (Blauer and others 1976; Stutz and others (1987) showed that diploid (2x = Collotzi 1966; Shaw 1992). It generally grows on well- 2n = 18) and tetraploid (2n = 36) populations differ drained soils of loamy to rocky or gravelly textures, in phenotype and distribution. In general, the dip- but it may occasionally be found on clay soils. Soils loids are restricted to the more central portion of usually range from neutral to alkaline; its adaptation the species range, are smaller (x = 13 inches tall and to salinity may vary ecotypically (McArthur and oth- 29 inches wide; 33.0 by 73.7 cm) than tetraploids (x = ers 1978b; Shaw1992; Stark 1966). It often grows in 16 inches tall and 37 inches wide; 40.6 to 94.0 cm) soils high in calcium (McArthur and others 1978b). and have narrower leaves (x = 3.8 mm versus x = 11.2 mm) than tetraploids (Stutz and others 1987). Spiny hopsage may occur in pure stands but more often is a component of communities dominated by other shrub species (Shaw 1992). It occurs in both Grayia spinosa ______sagebrush and salt desert shrub communities in association with Wyoming and basin big sagebrush, Spiny Hopsage blackbrush, black greasewood, shadscale, winterfat, Spiny hopsage differs markedly from spineless and pinyon and juniper species (Sampson and hopsage in many respects, but the fruits and flowers Jespersen 1963; Welsh 1984).

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Plant Culture received little use in that capacity. Glazebrook (1941), Wood and others (1976), and Kay and others (1977c) Seed production is highly variable between years, have all recommended its use for rehabilitation populations, and individual plants. During favorable plantings. However, additional work on site require- moisture years some shrubs produce large quanti- ments is needed to better assure artificial seeding ties of fruits, but during dry years few flowers or success. fruits are produced (Shaw 1992). Under warehouse conditions seed maintains viability up to 6 years Varieties and Ecotypes (Shaw 1992; Smith 1974a). Kay and others (1984) demonstrated that seed life can be extended by drying No improved varieties have been developed. Spiny seed at 35 ∞C (95.0 ∞F) for 6 days and storing them in hopsage is, however, composed of numerous ecotypes. glass jars with a desiccant at room temperature. This Use of these for various management purposes and procedure has maintained full seed viability for over 9 genetic improvement should both be possible. The years. species is known only at the tetraploid (4x) level Viable seeds germinate rapidly. For field estab- (Sanderson and Stutz 1994b). lishment late fall seeding is best because moisture levels are high (Shaw and others 1994). Bracts of spiny hopsage are probably important for seed dis- Kochia americana ______persal and seedling establishment (Shaw and others Gray Molly 1996). In some cases, however, removal of the bracts may be preferred for seeding depending on the equip- Gray molly is a small, woody-based subshrub to 20 ment used. Most drills are unable to seed the large inches tall (50.8 cm) (fig. 14). Its numerous, annual, fruits unless seeded alone. erect branches and leaves are covered with long, silky Young seedlings grow quite rapidly but become hairs. The linear, fleshy, more or less terete leaves dominant during the early summer. The plant often are 5 to 30 mm long. Hairy perfect or pistillate flowers grows without an understory of herbs, and the shrub are borne singly or in small clusters in the axils of seedlings are not highly competitive with herbs. To be leaves along almost the full length of the branches. successful, seedlings must be made in prepared seed- At anthesis, the hooded calyx lobes are about 1.5 mm beds free of competition. Small seedlings are vulner- long and closely cover the ovary. At maturity, the fruit able to weedy annuals. is largely concealed in the persistent calyx, which Broadcast seedings have erratic success. Natural develops conspicuous, horizontal, fanlike papery seeding is episodic; success depends on the availability wings to 3 mm long. This species blooms from June to of soil water (Shaw and others 1994). When hand August (Blauer and others 1976). seeded in pits or favorable sites and covered with soil, Gray molly usually occurs in saline or alkaline clay moderate success has been achieved. Rodents appear on plains and foothills between 4,500 to 6,000 ft to forage upon young seedlings, but the extent of their (1,400 to 1,800 m). Its range extends from southern damage is not fully understood. When seeded with Montana and eastern Oregon to New Mexico and herbs, small seedlings emerge and survive until about California. Gray molly summer cypress is quite widely midsummer. Seedlings then appear to become dormant, yet few recover the following season. Field grown transplants also demonstrate early summer dormancy even when irrigated. However, greenhouse reared stock do not senesce prematurely. Field survival of bare root transplants have been satisfactory when planted on mine disturbances and native sites. Neither container or bare root stock do well when planted offsite.

Uses and Management Spiny hopsage is a valuable forage for livestock and game from early May to midsummer while foliage persists. It receives little use during its dormant period (McArthur and others 1978b; Sampson and Jespersen 1963; Welsh 1984). Spiny hopsage furnishes good cover for upland game in all seasons. The Figure 14—Gray molly growing near Faust, plant has potential for use in revegetation but has Tooele County, UT.

488 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 20 Chenopod Shrubs used as winter forage by livestock when it is suffi- subspecies of forage kochia were introduced into the ciently abundant (Blauer and others 1976). It appears United States during the 1960’s (Keller and Bleak to have usefulness in revegetation of areas with a high 1974). The species is well adapted to the climate and salt concentration. soils of the Intermountain area. This is especially so for the pinyon-juniper, sagebrush, and salt desert Varieties and Ecotypes shrub communities (Blauer and others 1976; Keller and Bleak 1974; McArthur and others 1974; Stevens Both diploid (2x) and tetraploid (4x) populations and others 1985a). It grows at elevations in its native are known to exist but their characteristic differ- range from 0 to 8,000 ft (0 to 2,400 m) (Balyan 1972; ences, if any, are poorly understood (Sanderson and McArthur and others 1974). McArthur, n.d.; Sanderson and Stutz 1994b). Plant Culture Kochia prostrata ______Seed, under uncontrolled storage conditions, loses Forage Kochia viability rapidly in about 6 months (Balyan 1972; Jorgensen and Davis 1984). Seed viability can be main- Forage kochia is generally a long-lived, highly vari- tained up to 4 years by drying seed to 7 to 9 percent able, woody-based subshrub (fig. 15). However, some moisture levels and storing in airtight containers ecotypes may be definite upright shrubs. It ranges (Balyan 1972; Jorgensen and Davis 1984; Young and from less than 12 inches to over 40 inches (30.5 to others 1981c). 101.6 cm) in height. Ascending branches are covered In seeding forage kochia, best results occur when with short to long woolly hairs. Leaves are flat, linear the seed is surface sown and not covered (Stevens to filiform, and hairy. Flowers are borne in small and Van Epps 1984). Aerial seedings have been suc- clusters (glomerules) in the axils of slightly reduced cessful. Seeding at a rate of 0.25 lb (PLS) per acre leaves on the upper part of the stem. As the fruit (0.28 k/ha) is usually sufficient. If drilled, seeds should develops, the calyx forms dorsal appendages around be no deeper than 1⁄16 inch (1.6 mm) (Stevens and it that are rounded, flat, and tuberclelike, or oblong others 1985a). The species can be transplanted with and winglike (Shishkin 1936). Blooming occurs from good results in spring or fall when the ground is moist. July to September (Blauer and others 1976). Once established forage kochia is a good natural spreader. In concentrated stands, forage kochia at- Ecological Relationships and Distribution tracts large populations of lygus bugs, but such concentrations do not occur in mixed plant communities Forage kochia is native to arid and semiarid regions (Moore and others 1982). of central Eurasia extending to the Mediterranean basin and northeastern China. It grows on alkaline, Uses and Management stony, and sandy steppes and plains (Francois 1976; Shishkin 1936). Forage kochia tends to be evergreen. In its native habitat, forage kochia is a drought- Seeds of accessions from several types including both resistant, salt-tolerant species and is highly valued as a forage plant (Nechaeva and others 1977). The plant has high nutritive value with a high protein and carotene content (Balyan 1972; Davis 1979; Welch and Davis 1984). Forage kochia has its highest nutritive content in midsummer through fall when its crude protein and carotene levels are high (Davis 1979; Davis and Welch 1984). Frischknecht and Ferguson (1979), McKell and others (1979), and Ferguson and Frischknecht (1985) have shown forage kochia to be an effective mine spoil stabilizing plant. Sheep, goats, camels, and horses all consume it (Shishkin 1936). In Utah this species is sought out by mule deer. Davis and Welch (1985) showed some accessions are more palatable to mule deer than others. Forage kochia readily establishes from seed and grows rapidly, and under favorable conditions, reaches sexual maturity in 1 year (McArthur and others 1974). It spreads Figure 15—Forage kochia growing on a well, naturally, from seed. It is a useful component in roadcut near the mouth of Salina Canyon, shrub-grass pastures (Otsynia and others 1984). Once Sevier County, UT.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 489 Chapter 20 Chenopod Shrubs established, forage kochia will compete with annuals others (1996) recently confirmed the outstanding such as halogeton and cheatgrass (McArthur and performance of ‘Immigrant’ forage kochia in an adap- others 1990a; Monsen and Turnipseed 1990; Stevens tation and performance test across an environmental and McArthur 1990; Stevens and others 1985a). One gradient. Another accession (ssp. grisea) also per- desirable characteristic for competing with cheat- formed well. grass is its fire tolerance (McArthur and others 1990a). Forage kochia shows considerable potential for becoming a valuable forage and cover plant on arid Sarcobatus vermiculatus ______Western ranges. In arid shrublands, it is among the Black Greasewood most desired species to seed. Concern over possible poisonous properties is unnecessary (Davis 1979; Black greasewood is an erect, spiny-branched shrub Williams 1980). Russian workers have been seeding up to 10 ft (3.0 m) tall (fig. 16). The deciduous, bright- forage kochia on ranges since 1921 (Balyan 1972). green leaves 1 to 4 cm long are usually narrowly linear and often semi-terete. The shrub is usually monoecious. Ecotypes and Varieties Black greasewood had traditionally been considered as a member of the chenopod family. However, Although forage kochia is not native to North recent studies on chloroplast DNA and sieve element America, it is well adapted to many areas in Western plastids provide evidence that it should be separated North America (McArthur and Stevens 1983; Stevens from that family (Behnke 1997; Downie and others and others 1985a). A near relative, Kochia americana, 1997). Behnke (1997) reviews the taxonomic history is widespread but less robust and aggressive than of Sarcobatus and concludes that this genus is dis- K. prostrata (Blackwell and others 1978; Blauer and tinct enough to merit its own family, Sarobataceae. others 1976). We have included it here because of its traditional Forage kochia comprises two subspecies and two place in the Chenopodiaceae. varieties with separate habitat preferences (Balyan Ecological Relationship and Distribution 1972): Clay habitats; Kochia prostrata ssp. virescens Black greasewood grows from Alberta south into (green) Mexico with stands occurring in all the Western States. Sandy habitats; Kochia prostrata ssp. grisea (grey) var. A smaller species, Bailey greasewood (Sarcobatus canescens (densely baileyi), occurs in western Nevada in association pubescent) with shadscale and other desert shrubs. Shadscale Solonetzes (salty clay soils) var. villosocansa often grows with black greasewood as well. In the (white-villous) more saline areas of its range, black greasewood may be found in nearly pure stands. On such sites, Each of the above taxa has a relatively wide tolerance minor components of shadscale, Castle Valley clover to soil types; however, in general, the green subspe- saltbush, and Gardner saltbush may occur. In less cies (ssp. virescens) grows best in clay soils. It has saline areas, it may grow with threadleaf and whitestem both mountain and lowland ecotypes (Balyan 1972). Variety canescens of the grey subspecies (ssp. grisea) is adapted best to sandy habitats, whereas variety villosocansa is best adapted to salty clay soils (Balyan 1972). The species is a polyploid complex based on x = 9 with diploid, tetraploid, and hexaploid populations (Davis and Welch 1985; Herbel and others 1981; Pope and McArthur 1977). One selection, ‘Immigrant’, has been released (Stevens and others 1985a). This selection (ssp. virescens), originally from Stavrapol Botanical Gardens in Russia, has shown superior performance in longevity, forage production and quality, palatability, and competitiveness with annuals. It is adapted to sandy loam to heavy clay soils with best adaptation to heavier soils. ‘Immigrant’ has demonstrated its adaptability to the pinyon-juniper, basin and Wyoming big sagebrush, and greasewood- Figure 16—Black greasewood growing near Tuba shadscale shrub vegetative types. McArthur and City, Coconino County, AZ.

490 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 20 Chenopod Shrubs rubber rabbitbrushes, spiny hopsage, basin big sage- Varieties and Ecotypes brush, fourwing saltbush, winterfat, black sagebrush, and broom snakeweed. Greasewood often occurs in Black greasewood exhibits considerable variation valley bottoms where subsoil moisture is high. Often both in natural stands and plantations. Roos (1984), inland saltgrass, alkali sacaton, Nuttall alkaligrass, in a study involving 18 populations, concluded that alkali muhly, and basin wildrye grow with black Sarcobatus vermiculatus was rich in genetic varia- greasewood, they may form grassy understories. Rus- tion. Sanderson and others (1999) have documented sian wildrye, tall wheatgrass, and quackgrass have an interesting distribution of chromosome races in been successfully planted into black greasewood stands Sarcobatus. Sarcobatus vermiculatus, black grease- to make these stands more versatile grazing areas wood, is widely distributed in Western North America (McArthur and others 1978b). (fig. 1). Tetraploid (x = 9, 4x populations) occur only on the periphery of the distributional range to the north Plant Culture (southern Alberta, southern Manitoba, northern Mon- tana, eastern North Dakota), to the west (northeast- Like other chenopod shrubs, black greasewood pro- ern and east-central California and south-central Or- duces erratic seed crops that usually have low viabil- egon), and to the south (southern Arizona and ity. Seed is more difficult to collect because of the northwestern Sonora). Throughout the rest of the spiny, brittle nature of the stems. Poor seed quality species vast range including the central portion of that has decreased planting success. Seeds ripen in the range, S. vermiculatus is octoploid (8x). It is appar- fall and respond best to fall or winter seeding. Estab- ently an old plant; no diploids are known (Sanderson lishment by natural or artificial seeding is unpre- and Stutz 1994b; Sanderson and others 1999). Sarco- dictable. Seeding with grasses is not recommended batus baileyi is dodecapoid (12x) throughout its more unless the shrub can be seeded in separate rows. limited range (western and south-central Nevada) Black greasewood often exists on sites with clay- (Sanderson and Stutz 1994b; Sanderson and others textured surface soils. Soils compact and crust easily, 1999). often reducing seedling survival. Black greasewood has done well when seeded or transplanted on mine disturbances. Under these conditions it establishes Other Chenopod Shrubs ______well on fresh exposures and infertile soils. It competes well with herbs under these circumstances unless the Allenrolfea occidentalis (iodine bush), sites are fertilized heavily. Camphorosoma monspeliaca (Mediterranean camphorfume), Suaeda Uses and Management torreyana (desert sumpbush or desert Black greasewood contains soluble oxalates that blite) will sicken and kill hungry livestock that consume large amounts of the foliage (Kingsbury 1964). How- Iodine bush and desert sumpbush are two species ever, where there is a good assortment of other species, that grow in areas where salt concentrations are so this danger is negligible. It is often eaten by livestock high that they are the only forms of higher plant life (sheep and cattle) and wildlife in the spring. It is that survive (McArthur and others 1978b). In so usually not regarded as an important forage plant doing, they provide a valuable soil-binding service. although it furnishes cover and habitat. It is espe- Perhaps these species could be used more for reha- cially important on sites that flood, and it forms an bilitation work. Another species, an introduction important zone within various riparian habitats. It is from Eurasia, Mediterranean camphorfume, has highly important as cover to upland game birds and been shown to be useful in rehabilitating mine spoil big game animals. Sites that are burned, disked, or materials (Ferguson and Frischknecht 1985). This plowed and seeded to herbs receive heavy use. When species is similar in growth habit to forage kochia, seeded on mine and related disturbances, the shrub and like forage kochia, it is a useful forage plant in is often heavily grazed. south-central Asia (Nechaeva and others 1977).

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492 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 E. Durant McArthur Richard Stevens Chapter 21

Composite Shrubs

The sunflower family (Compositae or Asteraceae) is the largest family of flowering plants. Its many species occur around the world as annual and perennial herbs and as shrubs and trees (Benson 1957; Cronquist 1968; Wagenitz 1977). Three shrubby genera of the family—sagebrush (Artemisia), rabbitbrush (Chrysothamnus), and matchbrush (Gutierrezia)—make plants of this family among the most common and important plants of the Intermountain area (McArthur and others 1979a; table 1). Shrubs of these genera provide critically needed ground cover on arid Western ranges, are important sources of browse for domestic livestock and big game, and serve as cover and forage for many wildlife species. A number of sagebrush and rabbitbrush species are important as cover for small birds, game birds, and mammals, and as browse plants for big game animals, especially on winter and early spring ranges. Some species also provide forage for livestock (sheep and cattle). Horsebrush and matchbrush also contribute more forage than is generally believed; however, both plants may, under certain conditions, be harmful to domestic livestock (Benson and Darrow 1945; Johnson 1974a; McArthur and others 1979a) and cause allergies in humans (Lewis and Elvin-Lewis 1977; Rodriguez and others 1976).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 493 Chapter 21 Composite Shrubs

Chapter Contents by Species

General sagebrush culture ...... 496 Artemisia abrotanum (oldman wormwood) ...... 497 Artemisia arbuscula (low sagebrush ...... 499 Artemisia bigelovii (Bigelow sagebrush) ...... 500 Artemisia cana (silver sagebrush) ...... 502 Artemisia filifolia (sandsage or oldman sage) ...... 504 Artemisia frigida (fringed sage) ...... 505 Artemisia longiloba (alkali sagebrush)...... 506 Artemisia nova (black sagebrush) ...... 508 Artemisia pygmaea (pygmy sagebrush) ...... 509 Artemisia rigida (stiff or scabland sagebrush)...... 510 Artemisia spinescens (budsage)...... 511 Artemisia tridentata (big sagebrush) ...... 513 Artemisia tripartita (threetip sagebrush) ...... 521 Other sagebrushes ...... 522 General rabbitbrush culture ...... 522 Chrysothamnus depressus (dwarf rabbitbrush) ...... 524 Chrysothamnus linifolius (spreading rabbitbrush) ...... 524 Chrysothamnus nauseosus (rubber rabbitbrush)...... 525 Chrysothamnus parryi (Parry rabbitbrush) ...... 529 Chrysothamnus vaseyi (Vasey rabbitbrush) ...... 531 Chrysothamnus viscidiflorus (low rabbitbrush) ...... 531 Other rabbitbrushes ...... 533 Other composite shrubs...... 534 Haplopappus species (goldenweed) ...... 534 Tetradymia species (horsebrush) ...... 534 Gutierrezia species (matchbrush or snakeweed) ...... 534 Lepidospartum latisquamatum (scalebroom) ...... 534

Various forms of sagebrush and rabbitbrush may ranges include big sagebrush (A. tridentata), black also be directly seeded or transplanted for landscaping, sagebrush (A. nova), threetip sagebrush (A. tripartita), stabilizing, and beautifying disturbed landscapes and low sagebrush (A. arbuscula), budsage (A. spinescens), may have potential for supplying industrial chemi- fringed sage (A. frigida), sand sage (A. filifolia), cals. Young wildings of both sagebrush and rabbit- early sagebrush (A. longiloba), rubber rabbitbrush brush transplant easily. Usually within 3 to 7 years, (C. nauseosus), low rabbitbrush (C. viscidiflorus), direct seeded or transplanted plants are established Greene’s rabbitbrush (C. greenei), spreading rabbit- sufficiently to reproduce naturally from seed. Both brush (C. linifolius), and Parry’s rabbitbrush (C. parryi). establish well when properly aerially or drill seeded Some of these species, for example, threetip sagebrush (McArthur and others 1974; Plummer 1977; Plummer and Parry’s rabbitbrush, are usually found on ranges and others 1968). at higher elevations than the others. In this chapter, we prefer to limit the term sage- Virtually all of these species include an array of brush to members of the subgenus Tridentatae ecotypes. Some are large taxonomic units that include (McArthur 1979; McArthur and others 1981). Other several subspecies (big sagebrush, rubber rabbitbrush, woody members of the genus Artemisia then become low rabbitbrush, and Parry’s rabbitbrush) (McArthur “sage” or “wormwood.” However, in the broad sense in and others 1979a). These shrubs, especially under this chapter, use of sagebrush alone may refer to any conditions of heavy grazing, may form closed stands. woody, native North American Artemisia species. However, in many locations they mix with grasses and Sagebrush and rabbitbrush species that are espe- forbs. Big sagebrush often occurs with the grasses, cially important on low elevation Intermountain area bluebunch wheatgrass, western wheatgrass, the

494 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs

Table 1—Important Composite shrubs of the Intermountain burning, tillage, or mechanical or chemical treatment. area. Many of these observations were made on rangelands where the plant community was judged to be in near- Common name Scientific name climax condition. Some areas, in fact, had no evidence a of use or disturbance of any kind except by native Oldman wormwood Artemisia abrotanum fauna. On all study sites supporting subspecies of big Low sagebrush Artemisia arbuscula sagebrush, annual growth rings of the largest stems Bigelow sagebrush Artemisia bigelovii were counted to determine the age of the plants and Silver sagebrush Artemisia cana to establish the length of time during which there had Sand or oldman sage Artemisia filifolia been no fires. The oldest plants found were about 120 Fringed sage Artemisia frigida years old, and several locations supported plants at Longleaf sage Artemisia longifolia least 100 years old. Many of the older plants had split Alkali sagebrush Artemisia longiloba and nearly decayed stems on which the growth rings Black sagebrush Artemisia nova could not be accurately counted; so some plants may have been even older. With few exceptions, the oldest Birdsfoot sage Artemisia pedifida (not necessarily the largest) plants grew on stony or Pygmy sagebrush Artemisia pygmaea gravelly soils, and were of the Wyoming big sagebrush Stiff or scabland sagebrush Artemisia rigida subspecies. On disturbed relic areas, subspecies of big Budsage Artemisia spinescens sagebrush typically grew in open stands with indi- Big sagebrush Artemisia tridentata vidual plants or small clumps of plants uniformly Threetip sagebrush Artemisia tripartita distributed. Where it grew, big sagebrush made up five Alkali rabbitbrush Chrysothamnus albidus to 20 percent of the total annual production on the Dwarf rabbitbrush Chrysothamnus depressus study site. Spreading rabbitbrush Chrysothamnus linifolius Woody Artemisia, called wormwood in North Africa Greene’s rabbitbrush Chrysothamnus greenei and Eurasia, are principal components of the steppes Rubber rabbitbrush Chrysothamnus nauseosus of those regions. There (Larin 1956), as in North Parry rabbitbrush Chrysothamnus parryi America (Blaisdell and others 1982; Laycock 1979; Vasey rabbitbrush Chrysothamnus vaseyi Low rabbitbrush Chrysothamnus viscidiflorus McArthur and Plummer 1978), many range managers Small headed matchbrush Gutierrizia microcephala regard them as a mixed blessing. They are aggressive Broom snakeweed Gutierrizia sarothrae and increase on overgrazed land; on the other hand, Gray horsebrush Tetradymia canescens they provide much forage for wildlife and domestic Littleleaf horsebrush Tetradymia glabrata animals. This group of plants (Artemisia) may num- Nuttall horsebrush Tetradymia nuttallii ber 400 species and is divided into four subgenera Spiny horsebrush Tetradymia spinosa based on floral, anatomical, chemical, karyotypical, aIntroduced species. and distributional characteristics (McArthur 1979; McArthur 1983b; McArthur and others 1981; table 2). Rabbitbrushes are equally as widespread as sagebrush (= N. American Tridentatae) but are generally introduced crested wheatgrasses and desert wheat- less dominant (McArthur and Meyer 1987). There are grasses, and intermediate wheatgrasses, Idaho fes- 17 rabbitbrush species with 40 subspecies divided cue, sheep fescue, Indian ricegrass, Sandberg blue- into five sections (table 4). grass, Thurber needlegrass, bottlebrush squirreltail, Seed of most composite shrubs can be collected by and others. hand stripping, beating into a container, and, in some A rich array of forbs also grow in the sagebrush- cases, with vacuum harvesters (chapter 24). For seed grasslands, including species of aster, lupine, locoweed to pass through most seeders, plumes and other flower or milkvetch , balsamroot, Agroseris, wild buckwheat, parts need to be partially removed. This can be done by and goldenrod. Sagebrush species are the dominant passing the seed through a barley debearder followed and codominant species of over 40 described habitat by screening and fanning. Seeds are rather brittle; types of the American West (Blaisdell and others consequently, care must be taken to ensure that they 1982). Passey and others (1982) published a classic are not cracked or broken during cleaning and seed- study on sagebrush habitats of the Intermountain ing. Seed of most composites can be stored no more West. Their statement of the importance of sagebrush: than 1 to 3 years before seeding—otherwise, consider- Opinions differ among plant scientists as to the place able viability will be lost (Stevens and others 1981a). of sagebrush in plant communities on semiarid range- A principal trait of sagebrush and rabbitbrush is the lands (Ellison 1960). Historical accounts of the amount ability to persist under heavy abuse and poor manage- and distribution of sagebrush are also often contra- ment, and yet, be able to recover with proper manage- dictory. As noted, the relative amount and distribution of sagebrush species or subspecies have been ment. Few other shrubs can recover in a reasonable observed and evaluated on the several hundred tracts period without special management, including seed- examined in this study. One or more species of sage- ing. Most plants of this group can compete with weedy brush were present on every area examined except herbs, and many species occupy arid sites where few where there was unmistakable evidence of recent other shrubs persist.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 495 Chapter 21 Composite Shrubs

Important characteristics of a number of composite per lb (1.1 million per kg). In big sagebrush, seed size shrubs are listed in the Introduction to section VII. varies among subspecies. Basin big sagebrush has Seeding recommendations for principal vegetative just over 2.5 million seeds per lb (5.5 million per kg), types and conditions are discussed in chapter 17. Wyoming big sagebrush just under 2.5 million Composite shrubs adapted to these vegetative types (5.5 million per kg), and mountain big sagebrush has and conditions are included in the seeding recom- about 2 million (2.2 million/kg). Seed size, no doubt, mendations. Seed characteristics are found in chap- influences the depth at which seeds should be planted. ters 24 and 26. Seed of most sagebrush species should not be planted We include in this chapter a brief species descrip- deeper than 1⁄16 inch (1.6 mm). Good results can be tion, ecological relationships, distribution, culture obtained when seeds are broadcast on top of disturbed requirements, use, improved varieties, and manage- soils and not covered. ment of each of the species appearing in the chapter Adequate covering usually occurs from natural soil contents list. sluff. Sagebrush seed can be placed on top of the seedbed using drills that have had their seed drops or tubes pulled out between the disk-furrow opener and General Sagebrush Culture ______placed behind. Seeds then drop on the soil disturbed Intermountain sagebrush species all have small by the disk furrow openers. Several effective methods seed (see chapter 24). The smallest seeds are those of planting sagebrush seed include broadcasting (aerial of fringed sage, with over 4.5 million seeds per lb or by ground rigs), and placing through a seed drib- (9.9 million per kg) (100 percent purity). Pygmy sage- bler, thimble seeder, or browse seeder onto disturbed brush has relatively large seeds with about 500,000 soil. Drill seeding is a low priority option.

Table 2—Taxonomic subgenera of Artemisia (McArthur and others 1979a).

Classical Modern Distinguishing Species mentioned sectiona subgenerab characteristics Distribution in this chapter Absinthium Artemisia Pistillate ray flowers Eurasia A. abrotanum perfect disc flower, North Africa A. absinthium predominantly North America A. frigida herbaceous, but Abrotanum a few are woody A. longifolia A. ludoviciana Drancunculus Drancunculus Pistillate ray flowers Eurasia A. filifolia staminate disc flowers North America A. pedatifida herbacious and woody. A. spinescens Seriphidium Ray flowers lacking, Eurasia, Seriphidium perfect disc flowers; North Africad herbaceous and woody. Tridentatae Ray flowering lacking, North America A. arbuscula perfect disc flowers, A. argillosa woody.c A. bigelovii A. cana A. longiloba A. nova A. pygmaea A. rigida A. rothrockii A. tridentata A. tripartita

aDe Candolle 1837; Hooker 1840; McArthur and Plummer 1978; McArthur 1979. bRydberg 1916; Beetle 1960; Polyakova 1961; McArthur 1979; McArthur and others 1981. cThe single exception, A. bigelovii, has zero to two pistillate ray flowers on otherwise discoid heads. dTwo anomalous American species have been referred to Seriphidium: A. palmeri of southern and Baja California and A. mendozana Argentina.

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Seeding sagebrush with most ground equipment Artemisia abrotanum ______requires that the seed be cleaned to at least 60 percent purity. Most sagebrush seed is collected between 10 to Oldman Wormwood 15 percent purity. To increase purity, the collected material is run through a hammermill or barley Oldman wormwood, also called southernwood, is a debearder followed by fanning and screening. Barley shrub 1.5 to 6 ft (0.5 to 1.8 m) tall (Grieve 1931; Hall debearders are preferred (Booth and others 1997; and Clements 1923; Plummer 1974b). Stems are much Welch 1995). Hammermills tend to ball up the col- branched, erect or somewhat spreading, and form a lected materials, rendering it almost impossible to rounded bush (fig. 1). The species has a pleasing seed mechanically. aromatic fragrance and bright green, finely divided, As seed purity increases, volume of material de- pinnately dissected leaves giving the shrubs a feath- creases. Seed cleaned to more than 30 percent purity ery appearance. It has a deep and extensive root requires being mixed with a carrier such as rice hulls system allowing the plant to obtain nutrients and or screened sawdust. Range seeders and drills are not water from considerable distances (Plummer 1974b). equipped to handle seed the size of sagebrush seed. This characteristic also allows the plant to establish Small seed size requires that volume of material be in raw subsoils. Principal leaves are 3.0 to 6.1 cm long, increased to help ensure proper seed metering. Plant two or three times pinnately dissected with revolute materials and floral parts that are collected with margins, glabrous or sparsely puberulent on upper sagebrush seed will not flow through and will plug surfaces, but lightly tomentulose on the lower sur- most ground rig seeding devises. faces (Hall and Clements 1923). The inflorescence is Most sagebrush seed matures and is dispersed an elongated terminal panicle, leafy at the base, 6 to October through December. This is also the most ideal 15 inches (15.2 to 38.1 cm) long and 1.5 to 6 inches time to seed. We do not recommend spring seedings. (3.8 to 15.2 cm) broad. Heads are heterogamous, short Sagebrush seedlings are susceptible to frost damage and are intolerant of short drought. Many sagebrush seedlings that establish are located in, under, and next to downed and dead plant materials and next to rocks—all areas that provide some protection from adverse climatic conditions and grazing animals. When seeding sagebrush in a mixture with other species through ground equipment, we recommend that sagebrush seed be separated in the seeding device from seed of more competitive species (those with strong seedlings and fast rate of growth; most grasses fall in this group). Individual compartments in drill seed boxes can be constructed with cardboard partitions and duct tape. This will allow sagebrush and compatible species to be seeded separately but at the same time as seeding the noncompatible species another compartment. Seeding sagebrush aerially generally results in the seed of the various species in the mixture being separated so they are not in competition with each other. When dried and stored in an open, untreated, uncooled warehouse, seed of most sagebrush species will retain good viability for up to 3 years following collection (Stevens and others 1981a). Most sagebrushes transplant well. When standard bareroot and container stock handling, planting, site condition, requirements, principles, procedures, and methods are followed, excellent establishment success can be expected (Richardson and others 1986; Stevens and others 1981b). A majority of sagebrush seed is hand collected by Figure 1—Oldman wormwood plants growing stripping or beating the seed into some type of con- from stem cuttings along Ephraim Canyon Road, tainer. Seed of some species can be collected with a Sanpete County, UT. mechanical seed stripper.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 497 Chapter 21 Composite Shrubs peduncled, and nodding, and are about 2.0 to 2.5 mm Smaller cuttings root well but are difficult to stick in high, 2.5 to 3 mm broad. There are five to 15 ray the ground. For best results, leaves and small twigs flowers with 1.5 mm long corollas and 10 to 20 five- should be removed from the cuttings. Sharpening the toothed disk flowers with campanulate corollas, 1.5 to large end of the cutting will facilitate penetration of a 2 mm long. There are eight to 18 bracts. The receptacle hard soil surface and minimize damage to the cut- is naked. tings. The cuttings perform best if planted within 48 hours after being taken. However, cuttings have rooted Ecological Relationships and Distribution after 6 weeks in cold storage (32 to 40 ∞F; 0 to 4.4 ∞C). Cuttings should be kept moist and cool until planted. Oldman wormwood is native to the Mediterranean In general, roots are forming and new buds leafing out areas of Europe, Asia, and Africa (Hall and Clements about 2 weeks after planting. Plants should be placed 1923). Its alternate common name, southernwood, about every 2 to 4 ft (0.6 to 1.2 m) for good stands. distinguishes it from common wormwood (A. absin- Nursery plants remain vigorous and disease-free for thium). The latter is common and adapted in northern years (Nelson and Krebill 1981). Europe including Great Britain, whereas oldman wormwood was recognized as an introduction in these Uses and Management areas (Grieve 1931). In Britain, oldman wormwood rarely flowers, perhaps because of the high latitude. In Oldman wormwood is useful for stabilizing subsoil North America, the plant is occasionally adventive in sites (Plummer 1974b, 1977). Its deep extensive root the Eastern United States as a garden escape. In the system makes the plant especially useful for erosion Intermountain area, however, it occurs only where it control. Plants placed in staggered rows about 2 to 4 ft has been planted. Few viable seeds are produced and (0.6 to 1.2 m) apart make good ground cover. It is only rarely are seedlings found (Plummer 1977). A especially useful on steep sites such as roadcuts and dwarf form, var. nana 1 ft (30.5 cm) tall, and another fills. On such sites, new roots are produced on freshly regular sized ecotype have long been maintained by covered stems with each earth movement, thus pro- the USDA-ARS High Plains Research Station at Chey- viding additional stabilization. This species is useful enne, WY. Although seed production remains rare, a as a nurse crop in that it helps stabilize and modify raw soils so that natural invasion of plant materials from higher frequency of viable seed production has oc- the surrounding areas can occur. Individual plants curred with these additional plant genotypes growing may persist for 15 to 25 years or more, but eventually in proximity to the formerly near sterile ecotype. This they will go out without reproducing themselves. By suggests that outcrossing facilitates seed production that time, however, they have usually served their in oldman wormwood. It can be established by cuttings purpose. Plants are consumed by small animals (at in many soil types including raw subsoils in various times girdling the bark under snow) and insects, but vegetation types big sagebrush to the subalpine, 5,000 are not often eaten by large animals. Plants do provide to 10,500 ft (1,500 to 3,200 m) elevation, 12 to 40 inches excellent cover for upland game species. Formerly the (30.5 to 101.6 cm) precipitation (Plummer 1974b, plant was much used in herbal medicine as a tonic, 1976, 1977). It can tolerate moderately alkaline to anthelmintic, emmenagogue, and antiseptic (Grieve moderately acidic soils (Plummer 1974b). Although 1931). It is also used in ornamental plantings (Hall this species is adapted to subsoils such as roadcuts it and Clements 1923; Plummer 1974b). is not universally adapted to mine disturbances. This plant requires little management. Nurseries Plants are apparently sensitive to heavy metals or can be maintained as a source of cutting stock with other mine waste materials. Plants have not persisted only routine maintenance. Occasional weed control more than 5 years on acidic soils of the Idaho Batholith and supplemental water on dry sites is all that is (Monsen, unpublished). required. At outplanting sites where the plant has been put in to help with soil stabilization no care is Plant Culture necessary. These sites, however, often provide additional cutting stock. This species is, in essence, maintained vegetatively in the Intermountain area. It is easily propagated by Varieties and Ecotypes stem cuttings. The process for establishing the plant in a nursery or in outplantings is the same. Place Two varieties exist, the common one (var. abrotanum) cuttings in the ground in spring when and where soil and a dwarf one (var. nana). We know of no plant is moist. Plummer (1974b) gave instructions for cut- improvement work or ecotype comparisons. The com- tings. Stems 10 to 18 inches (25.4 to 45.7 cm) and up to mon variety is best, in most cases, for raw soil stabili- 0.5 inch (12.7 mm) in diameter give good results. zation planting because of its more vigorous growth.

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Artemisia arbuscula ______AB Low Sagebrush Low sagebrush (fig. 2) is a low spreading, irregularly branched shrub up to 20 inches (50.8 cm) high. The slender, erect twigs are densely canescent but may become nearly glabrous and thus darker green in late summer. The plant layers infrequently. Leaves are broadly cuneate, or fan shaped, 0.5 to 1.5 cm long, 0.3 to 1 cm wide, and are usually three-toothed (occasionally four to five in ephemeral leaves) or cleft at the apex (fig. 3). Leaves on the upper part of the flowering shoots may become entire. Flowerheads are grouped into elongated, narrow racemose panicles. The heads usually contain five to 11 disc flowers with corollas 3 to 4 mm long. The 10 to 15 involucral bracts are canescent. Flowering occurs August to September, depending upon strain and elevation. Seed ripens in October and November (McArthur and others 1979a).

Ecological Relationships and Distribution

Low sagebrush grows on dry, sterile, often rocky and Figure 3—Two subspecies of low sage- alkaline, usually clay, soils between 2,300 and brush: (A) A. arbuscula ssp. arbuscula; 11,500 ft (700 and 3,500 m) on approximately (B) A. arbuscula ssp. thermopola (Shultz 39,112 mi2 (101,300 km2) in 11 Western States (Beetle 1986). 1960; McArthur and Plummer 1978; Ward 1953). In the warmer, drier parts of its range, particularly in southwestern Utah and Nevada, it may grow well into and more rocky than those on which big sagebrush the mountains above 9,800 ft (3,000 m). In some areas, occurs. It may grow in mosaics with big sagebrush for example, east-central Idaho, low sagebrush occurs where each is confined to a particular soil type. Low on disjunct low and high elevation bands (McArthur sagebrush and black sagebrush only rarely occur in and others 1979a). Low sagebrush ranges from intermixed stands, for example, Lost River-Lemhi southern Colorado to western Montana and west Range area of Idaho (McArthur and others 1979a). In throughout Utah and Idaho to northern California, areas where the distribution of these two species Oregon, and Washington. Normally its sites are drier overlap, low sagebrush is usually found in the more moist habitats or at slightly higher elevations than black sagebrush (Tisdale and Hironaka 1981; Ward 1953; Zamora and Tueller 1973).

Plant Culture Plants can be established on adapted sites by direct seeding, broadcasting, and drilling. Seeding should be conducted in the fall on disturbed soil. Care must be taken to ensure that seeds are not covered more than 1⁄16 inch (1.6 mm). Wildings of this species can be transplanted in the spring or fall. Experimental nursery plantings have successfully reproduced. Plants of low sagebrush established at the Snow Field Station were more vigorous than most other sagebrush plantings there (Nelson and Krebill 1981). Plants establish well by direct seeding. Seedlings emerge rapidly and grow quickly, although few attain large Figure 2—Low sagebrush growing on shallow, stony soil in Logan Canyon, stature at first. Roots develop rapidly, and plants Cache County, UT. are able to survive dry surface conditions. Low

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 499 Chapter 21 Composite Shrubs sagebrush recovers quickly after burning and other others 1986; Winward and McArthur 1995). This disturbances. It persists and invades sites seeded to plant occurs in western Nevada. We believe that it is perennial grasses. It expresses good seedling vigor a stabilized hybrid between low and big sagebrush. and can be seeded with herbs. Seeds should be planted The big sagebrush parent is probably Wyoming big at a shallow depth with a firm seedbed. Seed lots of low sagebrush (Winward and McArthur 1995). sagebrush usually contain less inert or floral materials than processed seed lots of big sagebrush. Thus, seeds can be more easily planted. Seed of low sage- Artemisia bigelovii ______brush is more difficult to collect than the more robust Bigelow Sagebrush big sagebrush. Seed prices are, thus, somewhat higher, and lower quality seed is often sold. Low Bigelow sagebrush is a low shrub 8 to 16 inches sagebrush may often occur with little understory. (20 to 40 cm) high with numerous spreading branches When seeded with herbs, a mixed community nor- (fig. 4). It is also known as flat sagebrush (Hall and mally develops, but the understory often weakens Clements 1923). The flowering stems are slender and and recedes. The relationship with introduced and erect and bear inflorescences that are long, narrow native herbs is not well understood. panicles with short, recurved branches (fig. 5). New growth is covered with a silvery-canescent pubes- Uses and Management cence. The leaves of vegetative branches are similar to those of big sagebrush. They are narrowly cuneate, On winter ranges, and to a limited extent on summer 1 to 2 cm long, 2 to 5 mm wide, and normal tridentate, ranges, low sagebrush is browsed by big game and but may show various abnormal tips. The odor of livestock (Kufeld and others 1973; McArthur and crushed leaves is mild and pleasant. The heads are others 1979a). Considerable variation exists in how arranged into elongated, narrow panicles and nor- animals browse it in different locations. In Nevada a mally contain one, but occasionally zero to two, ray gray-green form may be heavily browsed, while the flowers and one to three, usually two, disc flowers. green form is only lightly browsed (Brunner 1972). In The turbinate involucre consists of eight to 12 short, addition to the direct browsing use of the plant, its densely tomentose, bracts 2 to 4 mm long and 1.5 to communities are important as habitat for a wide range 2.5 mm broad. Flowering occurs August to October. of domestic and wild animals (Dealy and others 1981). Bigelow sagebrush closely resembles, and is often Low sagebrush grows on difficult sites and is, there- mistaken for, low forms of big sagebrush produced by fore, not often a candidate for manipulation. Because overgrazing. In contrast to big sagebrush, it has ray low sagebrush grows on exposed sites it is important flowers. Furthermore, lobes of Bigelow sagebrush’s to wintering animals. Few other shrubs are adapted to vegetative leaves are always more shallow and more similar conditions. Consequently, maintenance of the sharply dentate than those of big sagebrush (McArthur species is often critical. Attempts to maintain low and others 1979a). sagebrush on sites not similar to its origin have proven unsuccessful. Seedlings usually establish when planted “offsite,” and plants may attain maturity. However, plants usually succumb after a few years with little or no regeneration.

Varieties and Ecotypes We are not aware of any substantial plant improvement work on this species. There are numerous ecotypes (Brunner 1972; McArthur and others 1979a) and three subspecies (fig. 3a,b shows two of these subspecies). Low sagebrush can apparently hybridize with its Tridentatae relatives (McArthur and others 1979a). Beetle (1959, 1960) named a dwarf form hotsprings sagebrush. This form occurs in the Stanley Basin area of Idaho, Jackson Hole area of Wyoming, east-central Oregon, and perhaps other locations. Beetle speculated that this form arose as a result of hybridization between typical low sagebrush and threetip sagebrush. Another important low Figure 4—Bigelow sagebrush growing at the Snow sagebrush is Lahonton low sagebrush (Winward and Field Station. The ruler is 12 inches (30 cm) long.

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Plant Culture Wildings can be successfully transplanted in the spring or fall. Nursery plantings successfully reproduce. Seedlings grow in the wild, suggesting that Bigelow sagebrush can be successfully direct seeded.

Uses and Management Bigelow sagebrush is palatable to livestock and game in all areas where it occurs. Its twigs are less woody, the odor milder, and taste less bitter than most of the big sagebrush complex (Hall and Clements 1923). This species can withstand considerable browsing; generally more than other sagebrushes. However, the plants are usually scattered, and consequently, not an abundant forage source except for some large stands in the Four Corners area.

Varieties and Ecotypes Ecotype variation has not been documented although it exists in chromosome races from 2x to 8x Figure 5—Bigelow sagebrush (McArthur and (McArthur and others 1981; McArthur and Sanderson others 1979a). 1999). Bigelow sagebrush occupies a taxonomic position between the true sagebrushes (subgenus Tridentatae) and the Artemisia (subgenus Artemisia) species. We have chosen to treat it as a member of the Tridentatae because of its characteristic growth habit, Ecological Relationships and Distribution woody anatomy, leaf form, chromosomal karyotype, and RAPD molecular genetic markers (McArthur and Bigelow sagebrush has a more southerly distribu- others 1981; McArthur and others 1998a). The confu- tion than other sagebrushes. It is one of the most sion arises because its flowers have zero to two ray drought-resistant of the sagebrushes. It occurs over 2 2 flowers as well as the characteristic disc flowers of the approximately 34,010 mi (88,086 km ) through west- Tridentatae. ern Texas, southern Colorado, New Mexico, Arizona, Utah, Nevada, and California in canyons, gravelly draws, and dry flats (fig. 6) from 3,000 to 7,900 ft (900 to 2,400 m) (Beetle 1960; Kearney and Peebles 1960; Ward 1953). This species is often found mixed with big sagebrush, black sagebrush, leafless green rabbitbrush, shadscale, and especially broom snakeweed. It is also characteristic in rocky soils of the southern portion of the pinyon-juniper woodlands (Hall and Clements 1923). Bigelow sagebrush is normally free of the common rust diseases and insect galls common on other Tridentatae taxa (Beetle 1960). However, in common with other Tridentatae, it is susceptible to a wilt disease (Nelson and Krebill 1981). Seedlings of Bigelow sagebrush spread well naturally. Seedlings grow rapidly and are robust and vigorous. Direct seeding of this shrub has been limited to experimental plantings. Where seeded, the shrub responds favorably and natural spread normally occurs around the parental plant. Plants appear to have good drought tolerance, even as young seedlings. Direct Figure 6—Bigelow sagebrush, foreground, seeding should not be a handicap to the use of this growing with fourwing saltbush in a gravelly species. draw near Bicknell, Wayne County, UT.

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Artemisia cana______Silver Sagebrush Silver sagebrush is an erect, freely branched, rounded shrub up to 5 ft (1.5 m) tall. Older branches have dark brown, fibrous bark while younger branches are covered with a dense white to yellowish-green tomentum. Leaves on the vegetative branches are 1 to 10 mm wide and 2 to 8 cm long, linear to linear-oblanceolate, entire or occasionally with one or two irregular teeth or lobes, silver-canescent becoming slightly viscid with age (fig. 7). Leaves on the flowering stems are similar, but they may be slightly smaller, especially on the upper parts of the stems. The foliage emits a mild to pungent aromatic odor when crushed. Numerous heads are arranged into dense, narrow, leafy panicles, some- Figure 7—Mountain silver sagebrush on Targhee times reduced to a raceme or spikelike inflorescence. National Forest, Sublette County, WY, showing Stems layer when in contact with the soil, thus pro- leaves and flower heads. ducing additional root systems. Each head contains four to 20 disc flowers. Ray flowers are lacking. Achenes are granuliferous. Blooming occurs during August and September. Seed ripens in October and November. success seeding silver sagebrush on mine disturbances on semiarid ranges in southern Idaho and Wyoming Ecological Relationships and Distribution and under nursery conditions. Seedlings develop rapidly often attaining a larger stature than plants of Aside from big sagebrush, silver sagebrush is the basin big sagebrush. Seedlings respond dramatically most widely distributed sagebrush. It occurs over to moist conditions and may reach 15 to 20 inches (38.1 2 2 approximately 53,221 mi (137,842 km ) in British to 50.8 cm) in 1 year. Plants are usually quite uniform Columbia and Saskatchewan in the north, south to in size and form. Young plants grow well with under- Arizona and New Mexico, and west to Oregon and story herbs. In undisturbed sites, conditions often California (Beetle 1960; McArthur and Plummer 1978). support a diverse understory. Plants appear well It grows on soils that are less mature, with less adapted to harsh disturbances when artificially seeded, phosphorus, potassium, nitrogen, organic matter, and but initially are slow to invade raw sites. lower cation exchange capabilities than do soils that support big sagebrush (Hazlett and Hoffman 1975). Use and Management However, it may grow on a variety of soils (Morris and others 1976). Silver sagebrush is important through its range as a browse shrub and is used quite extensively by live- Plant Culture stock and big game, particularly when other food is scare (Kufeld 1973; Kufeld and others 1973; Wasser Wildings of silver sagebrush can be transplanted in 1982). Bolander silver sagebrush was among the most the spring or fall. Establishment by direct seeding preferred of the sagebrush taxa offered to mule deer has also been successful (Kelsey 1986), especially and sheep in winter and fall feeding trials (Sheehy when done in the fall on the soil surface or at shallow and Winward 1981). Mountain silver sagebrush is depths (Wasser 1982). In most natural situations, grazed by sheep in the fall after grasses and forbs however, few seedlings survive (Walton and others dry. In the Western Great Plains, silver sagebrush is 1986). The species can also be propagated from hard- an important antelope survival food. Like big sage- wood cuttings. Harvey (1981) successfully rooted 87 brush, this species has been used by white settlers percent of the cuttings of plains silver sagebrush. and Native Americans for fuel (McArthur and others He used 6 inch (15.2 cm) cuttings with the basal, 1979a). Silver sagebrush is not as susceptible to fire 1.5 inch (3.8 cm) stripped of leaves, dipped in “rootone” as many other sagebrushes (Wright and others 1979; and placed in a misting propagation bench with a White and Currie 1983). In efforts to manage stand growth medium of 50 percent sand, 25 percent ver- densities by prescribed burning, White and Currie miculite, 15 percent peat, and 10 percent perlite. (1983) reported that on both fall and spring burns fire Monsen (personal communication) has had excellent intensity was directly related to plant mortality and

502 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs inversely related to subsequent growth. In general, layers whenever conditions are suitable. This sub- fall burns are more effective, as a silver sagebrush species may spread rapidly, particularly after burn- control measure, than are spring burns. Internal wa- ing, by rootsprouting and by rhizomes (Beetle 1960). ter stress in silver sagebrush is greater as the season Leaves of the vegetative branches are linear- progresses and soils dry out, and that stress is com- oblanceolate, entire or rarely with one or two irregu- pounded in thick stands where there is interplant lar teeth or lobes, 1 to 10 mm wide, 2 to 8 cm long, and competition (White and Currie 1984). An ecotype from are densely silky-canescent. Crushed foliage emits a the Sheridan, WY, area shows promise as a fire resis- pungent turpentine odor (Beetle 1960; Ward 1953). tant competition to cheatgrass (Monsen, unpublished). Flowerheads are usually arranged into dense, leafy panicles and may contain one to 20 disc flowers. Varieties and Ecotypes Blooming occurs during September, and the seeds ripen during October and November. It is octoploid, 8x No plant improvement work has been pursued on (McArthur and Sanderson 1999). Putative natural silver sagebrush. The species, however, hybridizes hybrids between plains silver sagebrush and big sage- with other sagebrushes (Beetle 1960; McArthur and brush subspecies have been found (Beetle 1960; Ward others 1979a; Ward 1953). It is, in all likelihood, a 1953). This subspecies has a more eastern distribution parent along with mountain big sagebrush for spicate than the other two silver sagebrush subspecies. It big sagebrush, which we believe to be a widely adapted occurs southern Canada southward, but mostly east of stabilized hybrid in Intermountain areas (Goodrich the Continental Divide, through Montana, the Dako- and others 1985; McArthur and Goodrich 1986). It tas, Wyoming, western Nebraska, and northern Colo- has also been proposed that silver sagebrush is one of rado. It grows particularly well on the well watered, the parents, along with early sagebrush, of the nar- deep soils of the northern Great Plains, especially rowly endemic coaltown sagebrush (Beetle 1960). Sil- along stream bottoms and drainageways (Johnson ver sagebrush is differentiated into three subspe- 1978; McArthur and others 1979a; Walton and others cies (fig. 8) separated from one another geographically, 1986). ecologically, and morphologically (Beetle 1960, 1977; Mountain silver sagebrush (A. cana ssp. viscidula) Harvey 1981; McArthur and others 1979a; Shultz is an erect shrub that readily layers. It usually is not 1983; Walton and others 1986). RAPD genetic mark- more than 3.3 ft (1.0 m) tall. Leaves on the vegetative ers confirm both the relationship between and the branches are 1 to 5 mm wide, up to 7 cm long, and are integrity of subspecies (McArthur and others 1998a,c). often crowded in dark green clusters. The leaves Plains silver sagebrush is an erect, round, canes- typically are simple and entire, but occasionally are cent, freely branched shrub up to 5 ft (1.5 m) tall. It variously toothed or lobed. This subspecies varies in appearance but is always darker green than mountain big sagebrush with which it is often growing (Beetle 1960). Mountain silver sagebrush is distinguished ABCfrom plains silver sagebrush by its smaller, darker green leaves, its lower stature, and more western distribution. Flowerheads are arranged into dense, short racemes or spikelike inflorescences 1 to 3 cm long. Each head contains four to 15 disc flowers. Flowers bloom during August and September. Seed matures during October and November. Mountain silver sagebrush occurs in mountainous regions around 5,500 ft (1,800 m) and above. It is usually found in areas of heavy, lingering snowpack from the southwestern corner of Montana, south along the Conti- nental Divide to New Mexico, and west to Arizona, Nevada, and Oregon (Beetle 1960, 1977; McArthur and others 1979a; Tisdale and Hironaka 1981; Winward 1980). It occurs as both diploid, 2x, and tetraploid, 4x, populations (McArthur and others 1981). Bolander silver sagebrush (A. cana ssp. bolanderi) has narrow leaves like mountain silver sagebrush, but they are canescent like plains silver sagebrush. It is Figure 8—Silver sagebrush subspecies: distributed in poorly drained, alkaline soils from central (A) A. cana ssp. cana; (B) A. cana ssp. Oregon to eastern California (Beetle 1960; McArthur bolanderi; (C) A. cana ssp. viscidula and others 1979a; Tisdale and Hironaka 1981; Winward (Shultz 1986). 1980). It is known only as diploid, 2x (Ward 1953).

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Artemisia filifolia ______Sandsage or Oldman Sage Sandsage is a round, freely branching shrub up to 5 ft (1.5 m) tall (fig. 9). Young branches are covered with a canescent pubescence while the older stems are covered by a dark-gray or blackish bark. Leaves are long, filiform, and silvery-white canescent; 3 to 8 cm long, less than 0.5 mm wide, entire or alternately divided into filiform divisions, and are often fascicled (fig. 10). Numerous, nodding heads containing two or three fertile, pistillate ray flowers and one to six perfect, but sterile, disc flowers are arranged into leafy, narrow panicles. Each head is subtended by five to nine canescent involucral bracts. Both the receptacle and achenes are glabrous. Flowers bloom during August and September. Seed ripens from October to December (McArthur and others 1979a). Other seed characteristics are given in tables in chapters 24 to 26.

Ecological Relationships and Distribution Sandsage is an excellent indicator of sand. It is probably the most widespread shrub on sand dunes and sandhills the southern Black Hills of South Dakota southward to Texas and Chihuahua and westward Figure 10—Sandsage (McArthur and others to Arizona and Nevada (Hall and Clements 1923; 1979a). McArthur and others 1979a; Rasmsussen and Brotherson 1986). In a study in southwestern Utah, Rasmussen and Brotherson (1986a) found that sandsage communities were floristically less diverse than adjacent plant communities, but that there was a higher density of plants in the sandsage communi- poor relative quality of the sandsage community soil ties. In the same study, they reported that despite the profile, sandsage nutrient quality was not affected. Sandsage accumulates mineral nutrients well above levels found in the soils in which it grows. At the study locations, at least, sandsage is adapted to soils of low fertility.

Plant Culture Wildings can be transplanted in the spring or fall. Plants can be easily reared from seed. Seeds are smaller than those of most other sagebrushes but grow well once established. Plants mature quickly even when planted offsite. Survival is reduced, however, when the species is planted out of its natural range (Nelson and Krebill 1981). It grows well on infertile soils. It has been seeded alone in most plantings but appears to develop rapidly and can be seeded with herbs adapted to arid sandy sites. It has been easy to grow as bareroot stock and survives well when field Figure 9—Sandsage growing near Moccasin, planted. It does not require special media for rearing Mohave County, AZ. (Monsen n.d.).

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Uses and Management Artemisia. Its range extends from Mexico northward through most of the western United States and West- The browse value of sandsage depends on where it ern Canada into Alaska, then on to Siberia, Mongolia, grows. It is seldom eaten in grasslands where other and Kazakhstan (Harvey 1981; USDA Forest Service food is adequate, but in more arid, desert regions it is 1937). Fringed sage is a common plant of the high consumed by cattle, sheep, and big game. It is particu- plains along the eastern slope of the Rocky Mountains, larly important in dry years (Hall and Clements 1923; but also occurs in valleys and mountains. In the McArthur and others 1979a). Sandsage is habitat for United States, it is most abundant in the eastern and some small nongame birds. This species helps pre- northern parts of its range. This species ranges from vent wind erosion by helping to stabilize light sandy low semidesert valleys to more than 11,000 ft (3,400 soils. It is an important candidate for harsh sandy m) elevation throughout the Rocky Mountain, Inter- sites but often grows in such arid sites that seed mountain, and adjacent plains regions. Fringed sage production is quite low. It is particularly useful in inhabits a wide variety of sites. Most typically, it blackbrush and pinyon-juniper sites where sandy out- grows in full sunlight in dry, coarse, shallow soils. On crops occur. winter ranges in western Utah and eastern Nevada, fringed sage may occur in dense stands along shallow depressions that collect moisture from summer rains. Varieties and Ecotypes In such areas, it is frequently associated with winterfat, No ecotypic differentiation has been documented, shadscale, and rabbitbrushes. On plains, foothills, and but it may well occur. It does have some cytological mountain slopes, this species may be associated with and chemical similarities with the true sagebrushes a variety of grasses and forbs as well as with various (subgenus Tridentatae) although it is assigned to the shrubs such as big sagebrush, Bigelow sagebrush, sandsage, and especially in overgrazed areas, with subgenus Dracunculus and differs from the Triden- broom snakeweed. It is a common understory plant in tatae in floral characteristics and wood anatomy ponderosa pine in several Western States (McArthur (Kelsey and Shafizadeh 1979; McArthur and Pope and others 1979a). 1979; Moss 1940).

Artemisia frigida ______Fringed Sage Fringed sage is a fragrant, aromatic, mat-forming perennial subshrub 8 to 24 inches (20.3 to 61.0 cm) tall (fig. 11). The lower woody stems are spreading and often much branched. Adventitious rooting may occur when stems contact the soil. The upper herbaceous stems are erect and leafy. The whole plant is densely silver-canescent. The numerous small, silky, canescent leaves are 6 to 12 mm long and are two or three times pinnately divided (fig. 12). This species has a deep perennial taproot with numerous extensive lat- erals that help it withstand drought. Numerous small flowerheads are borne in nodding racemes or open panicles. Small, densely hairy involucral bracts occur in several series around each flowerhead. Each head contains 10 to 17 outer, seed-producing, pistillate ray flowers and numerous (25 to 50) tubular, funnelform, perfect seed-producing disc flowers. Flower recep- tacles are densely villous. Fringed sage blooms from June at high elevations and latitudes, to November at lower elevations and latitudes. Seed matures between September and December (McArthur and others 1979a).

Ecological Relationships And Distribution

Fringed sage is probably, on a worldwide basis, the Figure 11—Fringed sage growing at Black most widely distributed and abundant species of Mesa, Gunnison County, CO.

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Plant Culture other areas, such as the grasslands of the Northwest and Great Plains, fringed sage may be less palatable Fringed sage can be established by seed or by and occasionally invades deteriorated grasslands. How- transplanting young plants or segments of old plants. ever, on the Great Plains, fringed sage is an important Surface seeding on disturbed soils is recommended winter antelope food and is used to a lesser extent the (Wasser 1982). It produces an abundance of small year round (McArthur and others 1979a). seeds, approximately 1,000 seeds per inch (2.5 cm) of It is well suited for growth with grasses and broad- inflorescence (Harvey 1981). There are 4.5 million leaf herbs. Its seedlings are competitive and can be seeds per lb (9.9 million per kg) (see chapter 24). In a direct seeded into areas with herbaceous competition. nursery, plants require little care and maintain rea- It is adapted to mine disturbances, perhaps better sonable vigor (Nelson and Krebill 1981). Individual than any other Artemisia. It can be used to control the plants are susceptible to overgrazing, but on a popula- rapid influx of weeds on large disturbances. tional basis, fringed sage seems to be stimulated and This species has strong reproductive qualities and is increases plant numbers with heavy grazing pressure a good pioneer shrub for stabilizing disturbed areas. It (Cooperrider and Bailey 1986). is often used in seeding western strip mines, especially coal areas. It has excellent reproduction by seed, and Uses and Management young plants or segments of old plants are readily transplanted in early spring (McArthur and others The forage value of fringed sage varies considerably 1979a). Its strong taproot and numerous lateral roots with location and season (Dietz 1972). Its value as make it effective in stabilizing gullies and reducing browse is highest in late fall, winter, and early spring soil erosion. These rooting characteristics make the on western ranges where it is eaten readily by big species capable of withstanding considerable grazing game and livestock (Cooperrider and Bailey 1986; and trampling use. Fringed sage has some value as a Kufeld 1973; USDA Forest Service 1937; Wasser 1982). medicinal plant (Hall and Clements 1923). It can be It is also important food for sage-grouse (Wallestad controlled when it is too abundant (Alley 1972). and others 1975). Nutritive quality is highest in the spring, but remains adequate for animals for much of the year (Cooperrider and Bailey 1986; Rauzi 1982). In Varieties and Ecotypes Undoubtedly, fringed sage, with its wide geographical and altitudinal distribution, has considerable genetic variation. However, no subspecific taxa are recognized and no plant improvement work is under way. Unlike several other widespread Artemi- sia species, fringed sage is known only at the diploid (2n = 2x = 18) chromosome level (McArthur and Pope 1979).

Artemisia longiloba ______Alkali Sagebrush Alkali sagebrush has also been called early sagebrush and low sagebrush. Alkali sagebrush is a mis- nomer because it grows mostly on neutral rather than on alkaline soils (Passey and Hugie 1962b; Tisdale and Hironaka 1981). It, however, is adapted to drought conditions (Robertson and others 1966; Zamora and Tueller 1973). Alkali sagebrush is a low shrub up to 18 inches (45.7 cm) tall (fig. 13). It has lax, spreading stems that frequently layer. The bark is dark brown to black on the older stems. The whole plant has a dark gray- green appearance (Beetle 1960). Leaves on the vegetative stems are broadly cuneate, up to 2 cm long, and are deeply three-lobed (fig. 14). Leaves of the flower- Figure 12—Fringed sage (McArthur and ing stems are similar but smaller. This species is others 1979a).

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Uses and Management The habitat of some stands of alkali sagebrush, when in good condition, supports a mixed understory of perennial grasses and annual and perennial forbs providing forage and cover for sage-grouse, antelope, and other wildlife as well as livestock (Dealy and others 1981; Monsen and Shaw 1986). Other stands are quite depauperate of associated vegetation and provide little forage and habitat for animals. Alkali sagebrush itself provides, depending on the site, browse for sheep (often preferred), and habitat and food for small animals (Dealy and others 1981; McArthur and others 1979a). The plant should be useful in rehabilitation plantings, particularly on heavy, seasonally dry soils. It has been observed in- Figure 13—Alkali sagebrush growing at vading roadcuts in Echo Canyon, UT, and barrow pits Wasatch Station, Summit County, UT. Note near Kemmerer, WY, (McArthur and others 1979a). the abundant, large flower heads. Monsen and Shaw (1986) reported that decadent stands can be rejuvenated by mechanical means. Sites that lack a satisfactory understory remain closed stands unless these shrubs are reduced in number by readily distinguished from other low statured sage- fire or mechanical means. brushes by its large heads and early blooming (Beetle 1959). Its heads contains six to 11 disc flowers and Varieties and Ecotypes are 3 to 5 mm broad as opposed to 3 mm or less for other short statured sagebrushes (low sagebrush and Alkali sagebrush has not been studied sufficiently black sagebrush). It flowers during mid-June to early enough to make substantial comments here. However, August and its seed ripens in August and September. because of its relatively broad and scattered distribu- Beetle (1960) points out that this species has in the tion and its differential use by animals, we suspect it past been confused with silver sagebrush because of harbors considerable genetic variation. Beetle (1960) its large heads; with big sagebrush because of its considered it to be a parent to the restricted endemic broadly cuneate, 3 lobed leaves; and with low sage- coaltown sagebrush. brush because of its dwarf size.

Ecological Relationships and Distributions Alkali sagebrush characteristically grows in heavy, highly impermeable soils generally with dense “B” horizons (Passey and Hugie 1962b; Tisdale and Hironaka 1981). It is, however, also found on lighter soils (McArthur and others 1979a). It occurs at elevations from 6,000 to 8,000 ft (1,800 to 2,400 m) along foothills and in basins of the ranges forming the Continental Divide in southwestern Montana, south through Wyoming to northwestern Colorado, and scattered westward through northern Utah and Idaho to Nevada and Oregon (Beetle 1960; Winward 1980).

Plant Culture Little is known except the plant can be transplanted. Natural seeding occurs rapidly following fires and other disturbances, yet mechanical tillage can limit seedling establishment if seeds are buried too deep or the seedbed is disrupted (Monsen and Shaw 1986). Figure 14—Alkali sagebrush (Shultz 1986).

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Artemisia nova ______Black Sagebrush Black sagebrush is a small spreading, aromatic shrub 6 to 8 inches (15.2 to 20.3 cm) or occasionally to 30 inches (76.2 cm) tall with a dull grayish-tomentose vestiture that causes most populations to appear darker than big sagebrush and low sagebrush (fig. 15). However, some forms may be as light in color as those species (Beetle 1960; Brunner 1972). Numerous erect branches arise from a spreading base (fig. 16). This shrub had not been observed to layer or stump sprout (Beetle 1960; Tisdale and Hironaka 1981). However, some layering of black sagebrush on a roadcut near Kolob Reservoir, Washington County, UT, has been noted (McArthur and others 1979a). Typical leaves are evergreen, cuneate, viscid from a glandular pubescence, 0.5 to 2 cm long, 2 to 8 mm wide, and three- toothed at the apex. The uppermost leaves, particularly on the flowering stems, may be entire. Flowerheads are grouped into tall, narrow, spikelike panicles that extend above the herbage. The inflorescence stalks are red-brown and persistent. The heads usually contain three to five disc flowers with corollas Figure 16—Black sagebrush (Shultz 1986). 1.8 to 3 mm long. The eight to 12 involucral bracts are greenish-yellow and nearly glabrous. The principal difference between black sagebrush and low sage- sagebrush, threetip sagebrush, and the subspecies of brush is that low sagebrush has five to 11 flowers per big sagebrush (Kelsey 1984). Three-fourths of a large head, 10 to 15 canescent involucral bracts, and is light sample (n = 152) of black sagebrush plants across the in color. Also, the flower stalks of black sagebrush are species range showed this characteristic conspicu- denser, much darker, and more persistent than those ously, whereas, no other taxa exceeded a 4 percent of low sagebrush (McArthur and others 1979a; Ward display (n = 1,849). 1953). Black sagebrush is characterized by the presence of leaf hairs readily visible at 10x magnification Ecological Relationships and Distribution when compared to its relatives low sagebrush, alkali Black sagebrush covers approximately 43,301 mi2 (112,150 km2) in the 11 Western States (Beetle 1960). It is most abundant at elevations from 4,900 to 7,900 ft (1,500 to 2,400 m) and normally grows on drier, more shallow stony soil than basin, mountain, or Wyoming big sagebrushes or low sagebrush (Beatley 1976; Tisdale and Hironaka 1981). Zamora and Tueller (1973) reported root restricting layers at depths of 11 to 27 inches (28.0 to 68.6 cm) in half of their black sagebrush study sites. Other soils in which black sagebrush commonly occurs are usually underlain by gravelly and sandy loam strata. Most, but not all, soils supporting black sagebrush are calcareous. Most black sagebrush stands do not burn because of their sparseness (Tisdale and Hironaka 1981). However, we have observed large burns of black sagebrush stands in central Utah. Insect galls are numerous on black sagebrush, but rust diseases are less common (Beetle Figure 15—Black sagebrush, with dark flower 1960). Nelson and Krebill (1981) found black sage- stalks, and Wyoming big sagebrush growing brush to be less susceptible to a wilt disease than at ecotone near Gabbs, Nye County, NV. most other Artemisia taxa growing at the Snow Field

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Station in Ephraim, UT. Clary (1986) reported that (Nagy and Regelin 1977); however, Smith (1950) and winter livestock grazing (principally sheep) has had a Sheehy and Winward (1981) found no such prefer- significant and often severe effect on black sagebrush. ence. Striby and others (1982) reported that mule deer Some large areas in Nevada formerly occupied by and elk preferred black sagebrush over basin big black sagebrush have been invaded recently by Utah sagebrush, but not over Wyoming and mountain big juniper and singleleaf pinyon pine. sagebrushes. However, Wambolt (1996) found that elk and mule deer preferred black sagebrush least in Plant Culture comparison with mountain, basin, and Wyoming big sagebrush. In the preference tests it should be re- Black sagebrush has been successfully transplanted membered that there are considerable intraspecific (Luke and Monsen 1984). Seed production in nurser- variations as well as interspecific ones being com- ies is often good. It should be seeded in fall or early pared. Black sagebrush has good winter nutritive winter. Seeds should be covered lightly up to 0.25 inch quality (Welch 1983b); not as good as big sagebrush, but (6.4 mm) deep. Once established, black sagebrush can the two are adapted to different habitats. Black sage- spread rapidly (Stevens 1986b). brush is a good conservation plant for dry, shallow, Seeds of black sagebrush are larger than those of stony soils. The plant is an aggressive natural spreader most sagebrushes (see chapters 20 and 24). Seeds from seed. Because it usually grows on dry rocky sites, tend to remain attached to the flower but can be proc- it is usually not a candidate for plant control. essed easier than seeds of other sagebrush. Seedlings are vigorous, and young plants often grow rapidly. Varieties and Ecotypes However, even under favorable conditions, young plants do not grow as rapidly as big sagebrush. Once Welsh and Goodrich (1995) recently described a new established, small seedings persist well even under variety, A. nova var. duchesnicola to go with the adverse conditions. Plants are able to establish amid longstanding typical taxon, A. nova ssp. or var. nova. rocky exposed surfaces. Under these conditions the Typical black sagebrush occurs in two color morphs shrub seedlings persist and usually exclude weedy (Beetle 1960). The gray-green form is generally browsed annuals. Black sagebrush can be established when more than the darker glossy-green form (Brunner seeded on sites other than its origin. It grows well on 1972; Stevens and McArthur 1974; Winward 1980). areas normally occupied by big sagebrush and may Welch and others (1981) and Behan and Welch (1985) persist and reproduce. Young stands grow well with have shown that an accession Pine Valley Ridge in seeded herbs, but as stands mature a reduction in Millard County, UT, is a fine candidate for wider understory herbs occurs. Black sagebrush establishes planting because of browsing animal preference. That well from direct seeding. The larger seed is not as accession has been transplanted and survived in sev- likely to be seeded too deep as smaller seeded species eral locations. It has been released as a germplasm of sagebrush. Under favorable moisture conditions, (Welch and others 1994). Black sagebrush has many mature plants produce abundant seed crops. Natural ecotypes that deserve additional attention. Black sage- spread occurs quickly, consequently interseeding or brush may have had a role in the parentage of Wyo- other techniques can be used to establish a seed source ming big sagebrush (McArthur 1983a; Winward 1975). and allow natural spread to populate large areas. Beetle and Johnson (1982) reported that some forms of black sagebrush approach Wyoming big sagebrush Use and Management in appearance. We have made similar observations on the Kaibab foothills in northern Arizona. Black sagebrush is usually held in high regard as a palatable forage for wildlife and livestock, especially Artemisia pygmaea ______sheep, antelope, and deer (Clary 1986; McArthur and Plummer 1978). However, it, like big sagebrush, has Pygmy Sagebrush populations that differ dramatically in mule deer preference (Behan and Welch 1985; Welch and others Pygmy sagebrush (fig. 17) is a dwarf, depressed, 1981). Welch and his colleagues (Behan and Welch evergreen, cushionlike shrub less than 8 inches 1985; Welch and others 1981) have shown that some (20.3 cm) tall. Bark on older stems becomes dark accessions are not eaten whereas others are highly brown and fibrous. On young branches the bark is preferred—one accession had 60 percent of its current nearly white to straw-colored and somewhat puberu- annual growth eaten in 1978 and 82 percent in 1982. lent. Leaves on the vegetative stems are green, nearly Scholl and others (1977) also reported accessional glabrous, viscidulous, 2 to 4 mm wide, 2 to 8 mm long, differences of mule deer preference between black and are pinnatified with three to 11 lobes, or some- sagebrush accessions. In comparison with other sage- times may be only toothed (fig. 18). Leaves on the brushes, black sagebrush was preferred by mule deer flowering branches are usually reduced and may be

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Utah to western Nevada and northern Arizona (Beetle 1960; McArthur and Plummer 1978; Ward 1953). In Nevada, this species is often associated with halophytic threadleaf rubber rabbitbrush. Some fairly large stands occur with black sagebrush in Utah. Because of its scarcity and small size, this species has little value as browse although it is eagerly taken when available. It does, however, provide important ground cover in the dry, alkaline areas where little else will grow. It establishes readily by transplanting divided plants. Although it spreads well from naturally dispersed seed, artificial planting of seed has not, as yet, been successful. This is due probably to poor quality seed and not the inability to establish by artificial planting. It is a good candidate to plant in heavy soils. It may be of considerable value for mine Figure 17—Pygmy sagebrush growing at the and roadway disturbances. Natural spreading has Snow Field Station, Sanpete County, UT. occurred when plants have been established in research plantings. entire. Heads with three to five disc flowers are ar- Artemisia rigida ______ranged into spikelike inflorescences. Ray flowers are lacking. Twelve to 18 greenish-yellow bracts subtend Stiff or Scabland Sagebrush each head. Achenes are glabrous. Flowers bloom in August and September, and seed matures in October. Stiff sagebrush is a low, pungently aromatic shrub Seeds are large for Artemisia (McArthur and others with thick, rigid, somewhat brittle branches up to 1979a; see chapter 24). 16 inches (40.6 cm) high (fig. 19). It is not known to Pygmy sagebrush is limited to calcareous soils in resprout or layer. The wider deciduous, silvery- desert areas over approximately 21 mi2 (54 km2) eastern canescent, spatulate leaves are mostly 1 to 4 cm long and deeply divided into three to five narrowly linear lobes (fig. 20). Occasionally some leaves are linear and entire. The inflorescence is a leafy spike with heads sessile or in small clusters in the axils of their subtending leaves, which generally are all longer than the heads. The campanulate involucre is 4 to 5 mm long with numerous, canescent bracts. Each head consists of five to 16 perfect disc flowers. Flowering occurs from late August to early October; seeds ripen in October and November (see chapters 20 and 24). This species resembles threetip sagebrush somewhat in its small size, silvery pubescence, and the deeply, narrowly lobed leaves, but may be distinguished by the spikelike inflorescence, large leafy bracts that subtend the heads, and the deciduous leaves. Stiff sagebrush occurs in dry rocky scablands in the Columbia and Snake River basins and spills over into the northern end of the Great Basin. It occurs primarily, if not exclusively, over basaltic bedrock (Daubenmire 1982; Tisdale and Hironaka 1981). It grows at elevations from 3,000 to 5,000 ft (910 to 1,500 m) in Idaho, central and eastern Oregon, and central and eastern Washington. Otherwise its ecological niche is similar to that of low sagebrush (Ward 1953). The report of this species growing in Montana are apparently in error (Morris and others 1976; McArthur and others 1979a). Figure 18—Pygmy sagebrush (Shultz 1986).

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Because of its scant foliage and stiff branches, stiff sagebrush has been considered of little value for browse, except for sheep (Hall and Clements 1923). It provides forage on the dry sites on which it grows in the midsummer when herbaceous plants are dry. This species provides cover on the poor rocky soils where it grows. It appears to have a wider range of adaptation than is indicated by its present natural range of occurrence, thus giving it potential use in reclamation of some harsh disturbed sites. Seed production is normally quite low and seed quality is questionable (Monsen, unpublished). At- tempts to seed and transplant this species in arid sagebrush ranges near Boise, ID, have not been successful. In most cases plantings were made on sites Figure 19—Stiff sagebrush growing at the Snow where cheatgrass and other annuals had invaded the Field Station, Sanpete County, UT. The ruler is shrublands. Stiff sagebrush seedlings were not able 30 cm long. to compete with the weeds. Plants grow slowly and appear to require 2 to 4 years to establish and attain reasonable size. The invasion of cheatgrass undoubtedly limits the natural distribution and occurrence of this shrub. However, it is an important plant for harsh rocky sites. Plantings have successfully established on prepared seedbeds. It is able to seed naturally on harsh sites and has been able to survive wildfires.

Artemisia spinescens ______Budsage Budsage is a low, spinescent, pungently aromatic, rounded shrub 4 to 20 inches (10.2 to 50.8 cm) high (fig. 21). It is profusely branched from the base and has white-tomentose pubescence on young twigs and leaves. This pubescence is grayish and stiff on older branches. Leaves are small, mostly 2 cm or less in length, including the petiole. Leaves are three to five palmately parted, with the divisions again divided into three linear-spatulate lobes. Leaves are crowded on the short stems, with those near the apex being smaller and more entire (fig. 22). Unlike most species of Artemisia, budsage is deciduous, with the leaves falling by midsummer. Early in the spring, when budsage first shows signs of breaking dormancy, but before the buds elongate, the bark the last season’s growth can easily be pulled off. At this developmental stage, budsage is sought out by big game and livestock. Sheepmen refer to this condition as “slipping.” As early as February or March, new bright-green leaves are produced. Budsage is well adapted to xeric conditions. It has an extensive root system that grows primarily in the top 6 to 22 inches (15.2 to 55.9 cm) of soil. Interxylary cork is formed annually over the last year’s wood in both the roots and the stem. This layer of cork restricts Figure 20—Stiff sagebrush (Shultz 1986).

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Terminal and lateral buds of budsage generally expand and begin to elongate in late March and early April during the latter part of the “slipping” period. Blooming normally occurs the last week in April through the last week in May, although it has been found in bloom as early as late March and as late as mid-June (Wood 1966). Although budsage ordinarily begins growth early in the spring and then becomes dormant by early or midsummer, it occasionally may break dormancy in response to late summer storms. The plants then remain green all winter to provide succulent forage throughout the winter and spring.

Ecological Relationships and Distribution

Figure 21—Budsage growing near Sevier Budsage is a drought-resistant shrub quite common Lake, Millard County, UT. in semiarid valley bottoms, benches, and foothills over much of the interior Western United States (Hall and Clements 1923; Wood and Brotherson 1986). It occurs the upward movement of water to the narrow zone of from Wyoming through southwestern Montana, wood formed by the current year’s growth. The corky southern Idaho, and eastern Oregon south to north- tissue develops during early summer and thus helps western New Mexico, northern Arizona, and south- to prevent excessive water loss during the dormant eastern California. It is often associated with shadscale, season (Wood 1966). Many other xerically adapted winterfat, and other salt-tolerant shrubs (McArthur woody Artemisia also have interxylary cork (Moss and others 1979a; Wood and Brotherson 1986). 1940). Budsage bears small flowerheads, 3 to 5 mm long, in glomerate racemes of one to three heads in leaf Plant Culture axils of the flower branches. Each head contains two to six fertile, pistillate ray flowers and five to 13 Summer dormancy can be broken by supplemental perfect, but sterile, disc flowers with aboritive ovaries. water. Wood and Brotherson (1986) reported that The loose flowerheads are held together by long, mat- supplemental water advanced the beginning of ted hairs that cover the corolla and especially the growth by 2 weeks, but did not increase total produc- achenes. The heads often fall off the plant intact, tivity by the time of the onset of cold weather. Seed without breaking apart to release the seed. However, quality is usually low, but during infrequent years, some seeds are usually dispersed independently. In natural seeding occurs. Some sites have improved well some instances, seeds germinate while still in the through protection from livestock grazing. Blaisdell head (Wood 1966). Seeds mature by early June. Good and Holmgren (1984) reported significant recovery of seed production occurs infrequently. The flowers budsage on areas protected from spring grazing. Arti- bloom so early in the spring that developing embryos ficial seeding has been hampered by the use of low frequently are frozen. Abundant reproduction occurs quality seed, not the inability of the plant to establish from direct seeding. Plants do not reach maturity in years of plentiful seed and favorable moisture. quickly, but seedlings and young plants are competitive. They persist under adverse conditions where few other species exist. Plants grow well with winterfat and scattered amounts of grass. Disking is detrimental to the survival of budsage, but anchor chaining usually has little effect upon the shrub.

Uses and Management Budsage is a palatable, nutritious forage plant for upland birds, small game, big game, and sheep in the winter. It is especially high in calcium, magnesium, phosphorus, and protein (Wood and Brotherson 1986). Generally, it is more palatable in the late winter than during the early winter (Holmgren and Hutchings 1972). During this time it is of tremendous value to Figure 22—Budsage (McArthur and others 1979a). the welfare of grazing animals, especially where there

512 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs is an abundance of dry grass. Care must be taken in grazing budsage in late winter and early spring. Even light grazing during this period is detrimental, and continual heavy grazing may eliminate budsage from areas (Holmgren and Hutchings 1972).

Varieties and Ecotypes Despite its wide distribution and isolated population, little morphological variation has been described in budsage. However, polyploidy is known (McArthur and Pope 1979).

Artemisia tridentata ______Big Sagebrush Big sagebrush (Artemisia tridentata) is a highly polymorphic species with numerous ecotypes and biotypes (fig. 23). Three subspecies—basin, Wyoming, and mountain (tridentata, wyomingensis, and vaseyana) big sagebrush—are generally recognized (Beetle 1960; Beetle and Young 1965) (see chapter 21). Additional subspecies, spicate big sagebrush (spiciformis) and xeric big sagebrush (xericensis), have recently been proposed (Goodrich and others 1985; Rosentreter and Kelsey 1991). Subspecies will be Figure 24—Large basin big sagebrush discussed in the Varieties and Ecotypes section. Big near Meeker, Rio Blanco County, CO. sagebrush is composed of aromatic, evergreen shrubs ranging from dwarf to tall, arborescent forms up to 15 ft (4.6 m) tall (fig. 24). The lower forms generally have several main stems arising from the base, whereas the tall forms often have a single short trunk. Older white to gray tomentum that gives the plants a silvery branches are covered with a gray to brown to black cast. Typical leaves are narrowly cuneate or oblan- shredded bark. Younger branches and leaves have a ceolate and terminate with three blunt teeth at their truncate apexes. However, considerable variation occurs, ranging from linear, entire leaves with rounded to acute apexes (fig. 25) to broadly cuneate leaves with varying numbers of teeth or shallow lobes. The leaves also range in size from 2 mm to 2 cm broad and 1 cm to 6.5 cm long. Normally, leaves of vegetative shoots are more characteristic and less variable than those on flowering shoots. Also, persistent, that is overwintering, leaves are less variable than leaves of the spring growth flush (Miller and Shultz 1987; Winward 1970, 1980), which are shed by midsummer. Heads of this species contain three to eight disc flowers, (except spicate big sagebrush, which may have up to 12 flowers per head) and are arranged into leafy panicles with erect or sometimes drooping branches. In some forms, the inflorescence becomes spikate. Blooming occurs from July to October. Seeds mature in October, November, and December. Big sagebrush plants often live 70 to 100 years or more. Specimens Figure 23—Individual big sagebrush plants of from several sites were found to be more than 200 basin, mountain, and Wyoming subspecies. years old (Ferguson 1964). Usually, however, a high Plants of each subspecies are randomly placed rate of individual plant turnover occurs within a 20 at the Gordon Creek Wildlife Management Area, year period (Stevens 1986b). Characteristics of big Carbon County, UT. Plants are even aged. sagebrush are detailed in table 3.

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Ecological Relationships and Distribution McArthur and Ott 1996). Nevertheless, his figures show how extensively sagebrush is distributed and Big sagebrush is the most widespread and common the relative abundance of each Artemisia species when shrub of Western North America. It is especially compared to related species. Big sagebrush grows in a common in the Great Basin. Beetle (1960) reports the variety of soils on arid plains, valleys, and foothills to species as covering approximately 226,374 mi2 2 mountain slopes from 1,600 to 11,200 ft (490 to 3,400 (586,309 km ) in the 11 Western States. Beetle’s esti- m) and is frequently associated with such shrubs as mate of coverage for this and other species is probably shadscale, saltbush, rubber rabbitbrush, low rabbit- too high, because it includes areas where Artemisia brush, fourwing saltbush, spiny hopsage, spiny are partial dominants (McArthur 1981, 1983b; horsebrush, winterfat, broom snakeweed, antelope bitterbrush, snowberry, and serviceberry. Although it is tolerant of quite alkaline as well as quite acidic soils, ABits optimum growth is in deep, fertile, alluvial loams (Sampson and Jesperson 1963). Although big sagebrush has spread with settlement of the West (Cottam 1961; Christensen and Johnson 1964; Hull and Hull 1974), it was clearly an important and widespread plant before this settlement. Vale (1975), quoting early pioneer and explorer diaries, has shown it was a common Western plant prior to 1850. Johnson (1986) presented photographic evidence showing sagebrush as a widespread dominant in the 1870’s. As early as the Pleistocene Epoch, sagebrush was already an important part of the Intermountain flora. Some 80,000 to 10,000 years ago, sagebrush dominated large tracts of land in areas where it is still found (Tidwell and others 1972; Van Devender 1977). Big sagebrush is an indicator of site quality (Kearney and others 1914; McArthur and Welch 1982; Passey and others 1982; Stevens and others 1974). Some populations, especially basin big sagebrush, have been taken out and the land they occupied put into agriculture. CDBig sagebrush and its subgenus Tridentatae relatives are subject to insect and microbial pests and benefactors—most notably the sagebrush defoliator moth (Aroga websteri) (see chapter 21). Wide ranging and periodic outbreaks of this insect have caused extensive sagebrush mortality over much of the range of A. tridentata and its relatives (Hall 1965; Henry 1961; Hsiao 1986; McArthur and others 1979a). How- ever, except on some critical winter game ranges, Aroga websteri is not believed to have serious, long- term effects. The moth is subject to insect parasites and predators and does not completely kill entire sagebrush stands (Hall 1965; Hsiao 1986). In time, sagebrush naturally reinvades its old sites that in the meantime have become more diverse plant communities. During drought, grasshoppers have been known to defoliate big sagebrush (Schlatterer, personal communication). Galls of many kinds of flies (Diptera) are found on sagebrush (Hall 1965; Jones 1971). The effect of the galls is not known. Sagebrush hosts many others insects of various orders (see Graham and Figure 25—Four subspecies of big sagebrush: (A) A. others 1995; Messina and others 1996). Some of tridentata ssp. tridentata; (B) A. tridentata ssp. spiciformis; these may protect the plant disease vectors. Several (C) A. tridentata ssp. vaseyana; (D) A. tridentata ssp. microbial-induced diseases are known (Krebill 1972). wyomingensis (Shultz 1986).

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Table 3—Characteristics of subspecies of Artemisia tridentata (adapted from McArthur 1983b).

Subspecies Characteristics tridentata vaseyana wyomingensis spiciformis References Habitat and range Foothills and Foothills and Foothills and valley High mountain Beetle and Young 1965; valley floors. mountains. floors. 2,500 to 7,000 ft areas. 7,000 to Morris and others 1976; 4,000 to 7,000 ft 3,000 to 19,000 ft (760 to 2,135 m). 12,000 ft (2,135 to Winward and Tisdale 1977; (1,220 to 2,135 m). (915 to 5,790 m). Montana, Washington 3,660 m). McArthur and Plummer 1978; British Columbia British Columbia to Arizona. Oregon and McArthur and others 1979a; and Montana to and Alberta to Montana to Dealy and others 1981; Baja California California and Nevada and Winward 1980; Harvey 1981. and New Mexico. New Mexico. Colorado. Smell Biter pungent Pleasant Bitter pungent Pungent, not bitter McArthur and others 1974. Essential oil x = 1.4 percent x = 2.2 percent x = 1.1 percent x = ? Welch and McArthur 1981. Leaf shape narrowly cuneate Cuneate to Cuneate Cuneate to Marchand and others 1966; spatulate narrowly McArthur and others 1974; cuneate McDonough and others 1975; Winward and Tisdale 1977. Common height 3 to 13 ft 2 to 5 ft 1.5 to 3 ft 2 to 5 ft McArthur and others 1979a; ranges (0.9 to 4 m) (0.6 to 1.5 m) (0.5 to 0.9 m) (0.6 to 1.5 m) Winward 1980. Sesquiterpenes 4 to 7 3 to 6 2 5 Kelsey and others 1973. compounds Ultraviolet Trace Abundant Trace, but often Abundant Shafizdeh and Melinknoff visible more than 1970; Stevens and McArthur coumarins tridentata. 1974; Brown and others 1975; McArthur and others 1981. Tendency to None Mild None Very strong Beetle and Young 1965; layer Winward 1980; Goodrich and others 1985. Palatability to Low Usually highest Highest for sheep High for sheep in Hanks and others 1973; deer and sheep sometimes high late summer and Sheehy and Winward 1981; for deer. fall. Sometimes Welch and others 1981; under snow in Striby and others 1982; season of use. Welch and McArthur 1986; unpublished; Welch and others 1987. Protein content High Low Low ? Welch and McArthur 1979b. Seed germination High Low Intermediate ? Harniss and McDonough prior to stratification 1976. 2n chromosome Commonly 18, Commonly 18, Consistently 36 18, 36 Ward 1953; Taylor and some 36 some 36 others 1964; Winward and Tisdale 1977; Kelsey and others 1975; McArthur and others 1981; McArthur and Sanderson 1999 Flower and seed Late Early Late-intermediate Early Marchand and others 1966; phenology Hanks and others 1973; Winward and Tisdale 1977. Flower stalk to x = 1.8 x = 3.4 x = 2.2 x = 3.1 Winward and Tisdale 1977. vegetative shoot length ratios Flower per head 3 to 6 4 to 8 3 to 8 6 to 12 Beetle and Young 1965; Winward and Tisdale 1977; Goodrich and others 1985. Effect of fire on Negative Positive Neutral Positive? Chaplin and Winward 1982. seed germination Leaf and stem water Lower Higher Lower Higher? Miller and others 1982; potential differences Shumar 1984; McArthur and others 1998b Shrub shape Uneven topped Even topped Uneven topped Even topped Winward 1980; Goodrich (usually) and others 1985. Position of Throughout Above crown Throughout Above crown Winward 1980; Goodrich flower stalks crown crown and others 1985. Seedling growth Longest sustained Slowest root Both above ground and ? Welch and Jacobson 1988; root growth rate growth rate root growth is initially Booth and others 1990. more rapid than other ssp.

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Some of these are widespread and may be locally Although established stands of big sagebrush can destructive; however, sagebrush populations are re- reduce the productivity of associated herbs, young silient and generally are not significantly affected in sagebrush plants do not diminish the density or vigor the long run. Nelson and Krebill (1981) isolated sev- of developing herbs. Consequently, sagebrush can be eral fungal species of the genera Gliocladium, directly seeded with herbs to provide a mixed compo- Fusarium, and Rhizoctonia from dying sagebrush in sition of plants. Big sagebrush is also compatible with uniform gardens. Similar symptoms (dying desiccated numerous shrubs, so mixed seedings can be done even plants) have been observed in natural populations though young sagebrush plants develop more rapidly (McArthur and others 1979a). Nelson and Sturges than serviceberry, snowberry, Martin ceanothus, (1986; Sturges and Nelson 1986) reported that green ephreda, and Wyeth eriogonum. snowmold disease can be serious in mountain big Big sagebrush wildings can easily be transplanted sagebrush populations. Diseases induced by these into nurseries and field locations when they are rela- fungi may be among the most serious sagebrush dis- tively dormant and moisture conditions are right— eases. Some microbes are likely useful for the vigor and growth of sagebrush. Wallace and Romney (1972) fall and early spring. Wildings about 6 to 8 inches found preliminary evidence that big sagebrush (15.2 to 20.3 cm) tall transplant best. McArthur and formed symbiotic relationships with microbial en- Plummer (1978) reported that of the 829 big sage- dophytes to fix atmospheric nitrogen. Williams and brush plants transplanted to the Snow Field Station Aldon (1976) found endomycorrhizae within big sage- in Ephraim, UT, in 1969 to 1970, 590 established brush roots and abundant spores around the roots. (71.2 percent). Five years later 421 plants were vigor- Endomycorrhizae, in general, have a beneficial influ- ous and growing. Luke and Monsen (1984) reported ence on plant growth by promoting nutrient absorp- high success in transplanting sagebrush on a Wyoming tion through infected roots (Williams and Aldon 1976). mine spoil site. Individual big sagebrush plants can For sagebrush, the beneficial effect has recently been be propagated by the rooting of stem cuttings demonstrated (Allen 1984). (Alvarez-Cordero and McKell 1979). Tissue culture propagation is also possible (Neville and McArthur Plant Culture 1986). Big sagebrush can be successfully drill (fig. 26, 27), Big sagebrush is one of few shrubs with potential for or broadcast seeded (fig. 28). Care should be taken that easy establishment by artificial seeding. Seedlings seeds are not covered too deeply, 1/8 inch (3.2 mm) or grow rapidly and compete well with herbaceous less is preferred (Monsen and Richardson 1984; plants. The plant establishes well from broadcast Richardson and others 1986; Young and Evans 1986b). planting on irregular but firm seedbeds. Consequently, Seeding should be done in the fall. Planting in subsoil plants can be established by broadcast seeding even substrates has had mixed success. Luke and Monsen when seeded in mixtures with herbs. Natural seeding (1984) and Monsen and Richardson (1984) were not often takes place amid established stands of seeded successful, but Stevens and others (1981b) success- grasses. Because natural spread generally occurs, fully seeded big sagebrush in subsoil scalps (fig. 29). sagebrush can be planted at low rates and allowed to spread. This has been successful. Sagebrush seed is difficult to separate from the chaff. Large stalks and debris are usually removed and seed is sold and seeded at a purity of 10 to 20 percent. Seeds are small and planting rates of 0.10 PLS (1 lb bulk per acre or 1.1 kg per ha) can produce adequate stands from broadcast or drill seeding. Seeds require minimal coverage but must be planted on a firm seedbed. Planting on a loose surface is not advised. Seeding should be delayed until early winter when soil surfaces are moist and firm. Seeds require afterripening and stratification to attain uniform germination. Small seedlings are not frost tolerant, and extensive losses occur from spring frosts. Sagebrush often establishes better from broadcast seeding than drill seeding, as seed are placed too deep with conventional drills. Seedings using the cultipacker Figure 26—Drill seeded big sagebrush with seeder are successful as seeds are not placed more grass mixture in southern Idaho. Sagebrush than 0.25 inch (6.4 mm) deep and the soil surface is seed was separated from grass seed in seed compacted or “firmed up” with this machine. boxes and drill rows.

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Figure 28—Plant establishment from aerial seeding of big sagebrush with crested wheatgrass, alfalfa, and Utah sweetvetch in south-central Utah.

reported that Wyoming big sagebrush was most palatable to deer and elk at their Montana study site. Wyoming big sagebrush is generally most palatable Figure 27—Seeded big sagebrush in subsoil to sheep (Welch and others 1987). Palatability of scalps in an intermediate wheatgrass seed- spicate big sagebrush has not been studied to any ing, Millard County, UT. great extent. It is browsed by sheep in the late summer and early fall and by big game when available in Matching seed source with planting site habitat is winter. Palatability of xeric big sagebrush is good for important for emergence and survival of big sage- mule deer and sheep. Sagebrush is important food and brush (Meyer and Monsen 1991). In general, regardless cover for upland birds. For example, sagebrush com- of subspecies, big sagebrush seed germination is prised 62 percent of the annual diet of sage-grouse in adapted to climate of collection sites (Meyer and oth- Montana (Wallestad and others 1975). Big sagebrush ers 1990b). A wilt disease can be troublesome in is an important component of antelope diet. Olsen and nurseries (Nelson and Krebill 1981). Symptoms of Hansen (1977) found sagebrush comprised 78 percent this disease are present also in natural populations. of the annual diet for antelope in Wyoming’s Red Snowmold can also cause lack of vigor and death in Desert. Even though sagebrush is a valuable forage, it sagebrush (Nelson and Sturgess 1986; Sturgess and is generally a less preferred browse for deer and other Nelson 1986). During cold winters with little snow cover, winter injury and death has been documented for large stands of sagebrush (Nelson and Tiernan 1983).

Uses and Management Big sagebrush is one of the more nutritious shrubs on Western winter livestock and game ranges. Basin big sagebrush has higher protein levels than the other subspecies (Welch and McArthur 1979b). Palatability of the different populations of this shrub to mule deer and other animals varies widely. For deer, basin big sagebrush is generally less palatable than Wyoming big sagebrush; both are generally less palatable than mountain big sagebrush (Hanks and others 1973; Scholl and others 1977; Sheehy and Winward 1981; Wambolt 1996; Welch and McArthur 1986; Welch Figure 29—An even-topped mountain big and others 1981). However, Striby and others (1982) sagebrush.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 517 Chapter 21 Composite Shrubs browsers than many rosaceous shrubs such as the the order and quantity of diversified associated spe- mountain mahoganies, bitterbrush, and cliffrose cies is vaseyana > tridentata > wyomingensis. This (McArthur and others 1978b; Smith and Hubbard information should be kept in mind for carrying capac- 1954; Smith and others 1965). Because of its wide- ity and manipulative considerations. spread abundance, its ability to grow with associated grasses, forbs, and other shrubs and its nutritious Varieties and Ecotypes nature, big sagebrush is the most important winter forage in foothill areas throughout much of the West Because of its plasticity and the apparent ease for livestock and big game. Sustained heavy use can, with which it hybridizes, there are good opportuni- however, result in loss of vigor and mortality of big ties for developing improved forms of big sagebrush sagebrush and other sagebrush species (Cook and for different purposes (McArthur 1981; McArthur and Child 1971; McArthur and others 1988a; Smith 1949). others 1979a, 1985, 1988b; Noller and McArthur 1986; It is one of the best shrubs available for use in reveg- Welch and McArthur 1979a). One accession has been etation of depleted winter game ranges in the Inter- released as a named germplasm; ‘Hobble Creek’ is a mountain area (Plummer 1974a; Welch 1983a). vigorous mountain big sagebrush from near the Meyer and colleagues showed that sagebrush seeds mouth of Hobble Creek Canyon, UT. It is palatable are site-adapted. Generally seeds from the area to be to mule deer and sheep; it persists in relatively dry planted or a similar area will establish and perform sites receiving from 13 to 14 inches (33.0 to 35.6 cm) of better (Meyer 1994; Meyer and Monsen 1992; Meyer rain; and has good growth characteristics (Welch and and others 1990b). others 1986). Other accessions also show promise and All forms of big sagebrush usually recover by natu- may qualify for later release. rally seeding sites that have been sprayed, burned, Basin big sagebrush (A. tridentata ssp. tridentata) disked, or chained. Natural spread can be reduced if is an erect, heavily branched, unevenly topped shrub soils are plowed or deeply disked causing seeds to be (fig. 24). Similar contrasting characteristics of basin deeply buried. Mountain big sagebrush recovers well and other big sagebrush subspecies are given in even amid the presence of established herbs. However, table 3. This subspecies has undivided or at least mountain big sagebrush usually grows on sites that trunklike, main stems. Most shrubs range between receive high amounts of summer moisture to sustain 3 to 6.5 ft (0.9 to 2.0 m) in height. Some forms, however, seedling growth. In addition, this sagebrush is able to may reach 15 ft (4.6 m) in suitable habitats. Mature produce good seed crops each year. Under more arid shrubs of this subspecies are the largest members of conditions, natural seed production is more erratic. the big sagebrush complex. The evergreen, vegetative Poor seed crops often occur and seed viability is dimin- leaves are narrowly lanceolate, up to 5 cm long by 5 mm ished after 2 to 3 years of storage (Stevens and others wide, and typically three-toothed at the apex. The 1981a). leaves of the flowering stems gradually become smaller Big sagebrush is aggressive and persistent and and may be linear or oblanceolate and entire. Winward sometimes forms closed stands. These stands may (1970) found the average length-to-width ratio of per- require thinning and rejuvenation. These techniques sistent leaves is 4.6 to 5.6. The gray-canescent foliage are treated in chapter 17, and in Plummer and others passes a strongly pungent, aromatic odor. Flowering (1955), Pechanec and others (1965), Laycock (1979), stems arise through the uneven crown and bear nu- Utah State University (1979) (especially Keller’s con- merous flowerheads in erect, leafy panicles. The heads tribution), Marion and others (1986), and Whisenant contain three to six small yellowish or brownish, (1986b). Big sagebrush stands are unexcelled in pro- trumpet-shaped, perfect disc flowers. The narrowly viding ground cover and forage when grazed to main- campanulate involucre consists of canescent bracts tain a balance between the sagebrush and associated 3 to 4 mm long and about 2 mm wide that form four to herbs and shrubs. Because big sagebrush has the five overlapping series around each head. The outer- potential to establish rapidly from both transplanting most bracts are less than a fourth as long as the and direct seeding, it is useful for stabilizing washes, innermost bracts. Flowering occurs from late August gullies, roadcuts, and other raw, exposed sites (Clary to October. Seed matures, depending on site, from and Tiedemann 1984; McArthur and others 1979a; October to early December. Seed can be collected fairly Monsen and Richardson 1984) and planting depleted easily by beating (see chapters 20 and 24). Basin big game ranges (Plummer and others 1968). Big sage- sagebrush is probably the most abundant shrub in brush, especially basin big sagebrush, has shown Western North America on lowland ranges. It nor- promise as a living snowfence (Laycock and Shoop mally occurs on dry, deep, well-drained soils on plains, 1986). Winward (1980, 1983) reported that associated valleys, and foothills below 7,000 ft (2,100 m) elevation. species richness with each subspecies (see next sec- Above this elevation subspecies vaseyana and spici- tion) varies greatly. Of the three common subspecies, formis, and occasionally subspecies wyomingensis are

518 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs more prevalent. Vigorously growing basin big sage- adapted to dry sites between 2,600 and 4,600 ft (790 to brush is considered indicative of productive ranges 1,400 m) in southern Idaho and surrounding areas. Its because it often grows in deep, fertile soil. This sub- introgression with ssp. tridentata is apparent in that species has generally been regarded as intolerant of plants are large and ragged topped, but chromato- alkali, but there are ecotypes that grow in relatively graphically, cytologically, and phenologically it most high alkalinity in association with such alkali-tolerant closely resembles ssp. vaseyana. plants as black greasewood, shadscale, and saltgrass. Wyoming big sagebrush (A. t. ssp. wyomingensis) is A similar taxa restricted to southern California is ssp. somewhat intermediate in distribution, ecology, and parishii distinguished by its exceptionally hairy morphology between basin big sagebrush and moun- achenes. Subspecies parishii may only be an ecotype of tain big sagebrush, but more diminutive than either basin big sagebrush. (fig. 30). It is adapted to drier sites than other subspe- Mountain big sagebrush (A. t. ssp. vaseyana) is nor- cies by virtue of its rapid and long root growth and mally a smaller shrub than basin big sagebrush. Its early, but relatively small, aboveground growth (Booth main stems are usually divided at or near the ground, and others 1990; Welch and Jacobson 1988). The and it tends to have a spreading, evenly topped accession ‘Gordon Creek’ has recently been released as crown (fig. 29). Vegetative branches are usually less a named germplasm (Welch and others 1992b). It than 3 ft (91.4 cm) high and occasionally layer at the combines wide adaptability, palatability, favorable base. Some lower elevation ecotypes reach about 6 ft growth characteristics, and drought tolerance. Occa- (1.8 m) in height. The persistent vegetative leaves are sionally, the subspecies tridentata, vaseyana, and broadly cuneate to spatulate and are characteristi- wyomingensis may be found growing together. cally wider than those of both basin and Wyoming big Whenever it is found associated with ssp. tridentata, sagebrushes. When looking down at shrubs of moun- ssp. wyomingensis is growing in the poorer, more tain big sagebrush, terminal leaves on each twig shallow soils (Beetle and Young 1965). Subspecies appear to be distinctly whorled. Basin big sagebrush wyomingensis is a low shrub usually less than 3 ft does not show this trait, but Wyoming big sagebrush (91.4 cm) in height. It has an uneven to round top with shows the trait to some extent. Mountain big sage- flower stalks arising throughout the crown like ssp. brush grows in slightly acidic to slightly alkaline soils tridentata. Its main stem branches at or near the (Welch and McArthur n.d.). Unlike ssp. tridentata, ground level like ssp. vaseyana, but it does not layer. ssp. vaseyana is rarely associated with any of the Leaves are 1 to 2 cm long, narrowly cuneate to cuneate, saltbushes. Mountain big sagebrush has larger seeds and have an average length-to-width ratio of about 3:1 than Wyoming and basin big sagebrushes. It usually for the persistent leaves (Winward 1970). Flowerheads comes better from direct seeding than the other two contain three to eight disc flowers and are arranged subspecies. There are two varieties of mountain big into panicles narrower than the paniculate inflores- sagebrush. Each variety occurs in large distinct cence of tridentata, and is wider than the spicate inflo- populations but also integrates into other populations. rescence of vaseyana (Beetle and Young 1965; Winward The most common variety outside of the Pacific North- and Tisdale 1977). Flowering and seed ripening take west is small-headed mountain big sagebrush (Goodrich and others 1985). This is the sagebrush that Ward (1953) referred to as that of “timbered or mountainous areas in which the plants are very uniform in size, usually about 2 ft (61.0 cm) in height, and of a rather spreading, flat-topped habit of growth, with the inflorescence extending upward like plumes above the rest of the bush.” The small-headed form has four to six flowers per head. The other variety, var. vaseyana, has seven to 11 flowers per head and has a more restricted geographical range in the upper elevational sagebrush areas of Washington, Oregon, and Idaho with only occasional small patches occurring outside of that area (Goodrich and others 1985). Rosentreter and Kelsey (1991), on Winward’s (1970) suggestion maintained that xeric big sagebrush, (A. t. ssp. xericensis) be recognized as a distinct entity. This subspecies is apparently derived from hybridization between ssp. tridentata and vaseyana (McArthur and Figure 30—Wyoming big sagebrush growing others 1979a; Rosentreter and Kelsey 1991). It is near Daniel, Sublette County, WY.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 519 Chapter 21 Composite Shrubs place later than vaseyana and earlier than tridentata. Wyoming big sagebrush plants have an effective near surface root system (Sturges 1977). This may be one reason why relatively few plant species grow with it. This subspecies may have arisen from hybridization between ssp. tridentata and vaseyana (Beetle and Young 1965; Hanks and others 1973) or ssp. tridentata and A. nova (Winward 1975), or a combination of all three taxa (McArthur 1983b). Wyoming big sagebrush grows throughout the Intermountain region. It is particularly abundant east of the Continental Divide in Montana, Wyoming, and parts of Colorado in dry, shallow, gravelly soil; usually from 5,000 to 7,000 ft (1,500 to 2,100 m) (Beetle and Young 1965). In Idaho, this subspecies is found from 2,500 to 6,500 ft (760 to 2,000 m) in the hotter, drier portions of the state (Winward 1970). Timberline big sagebrush (A. t.ssp. spiciformis) has been the subject of some taxonomic confusion (see A. rothrockii discussion in Goodrich and others 1985 and McArthur and others 1979a, 1981). This taxon is Figure 32—A specimen of timberline big sagebrush collected from Wolf Creek Summit, Wasatch the widespread root sprouting big sagebrush of high County, UT. elevations in the Intermountain area (Goodrich and others 1985; McArthur and Goodrich 1986). Its habitat is often in openings in aspen, spruce, or fir woods or where drifting snow accumulates (fig. 31). It had big sagebrush in stem and leaf characteristics, but formerly been confounded with Rothrock sagebrush mountain silver sagebrush in floral characteristics (Artemisia rothrockii), a similar taxon of the Sierra (fig. 32). It has 10 to 18 flowers per head. Timberline Nevada (Goodrich and others 1985; Shultz 1983; Ward big sagebrush flowers in August and September in 1953). Rothrock sagebrush differs from timberline native habitats, but in early June when transplanted sagebrush by its dark resinous leaves, restricted to low elevations (4,500 to 5,600 ft; 1,400 to 1,700 m). California distribution, and high polyploidy. Timber- It is a palatable sagebrush often browsed heavily in line big sagebrush is thought to be a stabilized hybrid the fall by sheep and mule deer. In winter it is usually between mountain big sagebrush and mountain silver unavailable under snow. It has shown potential as a sagebrush (Goodrich and others 1985). In growth low landscape hedge (McArthur and others 1979a). form, timberline big sagebrush resembles mountain Although basin, Wyoming, and mountain big sagebrush occur in distinct sites, all three of these common subspecies express a wide range of adaptation. When planted throughout the big sagebrush, pinyon- juniper, and mountain brush communities all three subspecies demonstrate adaptiveness. Although sources from high elevations are not suited to arid circumstances, and sources collected from light textured soils are not adapted to heavy clay textured soils, most selections have established and persisted when planted over a wide range of sites. The subspecies of big sagebrush as well as near relatives (=subgenus Tridentatae) hybridize at many of their contact points. In fact, hybridization is thought to have been a key mechanism in the evolution and speciation of the group (Beetle 1960; Hanks and others 1973; McArthur and others 1981, 1988; Ward 1953). Artificial hybridization may be useful in selecting and Figure 31—Timberline big sagebrush growing combining traits for management purposes such as near Minturn, Eagle County, CO. Note the fire tolerance and palatability (McArthur and others large flower heads and flowering stalks. 1979a, 1988b,c, 1998a; Weber and others 1994). Some

520 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs current studies of big sagebrush zones of hybridiza- AB tion are yielding results in an effort to better under- stand the role of hybridization in nature and to help determine the importance of the permanence and function of hybrid zones. These studies have concentrated on morphological, chemical, physiological, cytological, ecological, edaphic, and entomological characteristics (Byrd 1997; Byrd and others 1999; Freeman and others 1991, 1995, 1999; Graham and others 1995; McArthur and others 1988b,c, 1998a, 1999b; McArthur and Sanderson 1999; Messina and others 1996; Wang 1996; Wang and others 1997, 1998, 1999).

Artemisia tripartita ______Threetip Sagebrush Threetip sagebrush is a round, evergreen shrub up to 6 ft (1.8 m) high. It may have a simple, trunklike main stem or several branches arising from the base. The bark on young branches is canescent but becomes Figure 33—Threetip sagebrush (Shultz 1986): shredded and grayish, light brown to dark brown or (A) A. tripartita ssp. rupicola; (B) A. tripartita black on older stems. This species can layer, some- ssp. tripartita. times sprout back after a burn, and may sprout from the stump following herbicide treatment (Beetle 1960; Pechanec and others 1965; Schlatterer 1973; Winward 1980). Leaves of the vegetative branches are canes- Uses and Management cent, 0.5 to 4 cm long, and typically deeply divided into three linear or narrowly linear-lanceolate lobes, Threetip sagebrush is a vigorous seeder, but unfor- which in turn may be three-cleft (fig. 33). Some of the tunately some forms are not particularly palatable. upper leaves are often entire. Crushed foliage emits a However, Kufeld (1973) reported use by elk, and Kufeld pungent odor. Flowerheads contain three to 11 disc and others (1973) reported use by deer. Beetle (1960) flowers and are normally arranged into panicles. Ray reported it was of low palatability for both livestock flowers are lacking. Each head is subtended by eight and game, and Brunner (1972) observed that it was to 12 canescent involucral bracts. Achenes are resin- never grazed in Nevada. However, a form near Salmon, ous-granuliferous. Blooming occurs from July to ID, is palatable to deer. The form near Salmon may have been introgressed by a nearby population of September and seed is collected from mid-October to Wyoming big sagebrush (McArthur and others 1979a). mid-December. Stands of threetip sagebrush sometimes require thinning by mechanical and chemical means Ecological Relationships and Distribution (Pechanec and others 1965). Threetip sagebrush covers approximately 13,000 mi2 Threetip sagebrush has not been difficult to estab- 2 lish by direct seeding, rearing, or transplanting. Usu- (33,670 km ) in the Northern Rocky Mountains and ally a large amount of high quality seed is produced Great Basin States from British Columbia south annually. Seeds can be harvested without difficulty through Montana and Wyoming to Colorado and west from the upright bushes. Within its area of occurrence, to Washington, Oregon, northern Nevada, and north- threetip sagebrush establishes and performs well. ern Utah at elevations between 3,000 to 9,000 ft Seedlings are able to persist within both a dense (910 to 2,700 m) (Beetle 1960). In some places, particu- perennial and annual understory. But threetip sage- larly in Idaho, this species occurs between the lower, brush is not any more competitive with cheatgrass hot, dry sites dominated by Wyoming big sagebrush and other annuals than other species of sagebrush. and the higher, cooler sites dominated by mountain Plants grow quickly and can attain a mature stature big sagebrush (Schlatterer 1973). It usually grows on in 3 to 5 years. The shrub frequently exists with a moderate to deep, well-drained, loamy and sandy loam number of other shrubs and herbs. Seeding a mixture soils (Winward 1980). of species usually does not reduce the survival or productivity of this shrub.

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Attempts to establish threetip sagebrush onto arid Other Sagebrushes ______sites dominated by Wyoming big sagebrush have not been successful. Plants initially appear but fail to Coaltown sagebrush (Artemisia argillosa) is a three- persist if reseeded. However, some ecotypes can be tip form endemic to a small area in Jackson County, extended to drier sites. CO. This species is well adapted to heavy clay spoil Threetip sagebrush recovers well following exten- material and may have some utility in reclamation sive disturbances. However, hot summer fires can kill activities (Beetle 1960; McArthur and others 1981). large stands and prevent resprouting and can signifi- Rothrock sagebrush (A. rothrockii) is similar in cantly delay recovery by seeds. many respects to subalpine big sagebrush and would Threetip sagebrush often grows intermixed, but share the latter’s uses. However, Rothrock sagebrush as separate stands, with mountain big sagebrush, does not occur in the area of interest for this book. It is Wyoming big sagebrush, or alkali sagebrush. When confined to the high country of California (Goodrich disturbances have occurred in these areas, threetip and others 1985; Shultz 1983). sagebrush seedlings often invade into all circum- Two other species—longleaf sage (A. longifolia) stances. Natural sorting occurs, but rather slowly. and birdsfoot sage (A. pedatifida)—are not true sage- Threetip sagebrush demonstrates good compatibility brushes in that they don’t belong to the subgenus with many introduced and native grasses and broad- Tridentatae (table 2). Both, however, may prove useful leaf herbs. It survives and grows well with competitive in revegetation efforts. Longleaf sage is the most grasses including mountain brome, slender wheat- robust and woody member of the mostly herbaceous grass, intermediate wheatgrass, and orchardgrass. Louisiana sage (A. ludoviciana) complex (see chapter 19 for treatment of the Louisiana sage or sagewort Varieties and Ecotypes complex). Longleaf sage may grow up to 3 ft (91.4 cm) tall. It grows well on clay soils from Nebraska and Wyoming threetip sagebrush (Artemisia tripartita Wyoming north to Canada (Hall and Clements 1923). ssp. rupicola) is a dwarf shrub with decumbent Birdsfoot sage is a low perennial subshrub up to 6 branches that rarely grows over 6 inches (15.2 cm) tall. inches (15.2 cm) tall with a tough woody root. It grows It is frequently found layering and may have a crown on alkaline flats in Wyoming and Idaho and could be of 12 to 20 inches (30.5 to 50.8 cm) (Beetle 1960). used for rehabilitation of alkaline spoils (Hall and Leaves of the vegetative branches are often 3 cm long Clements 1923). and deeply divided into linear lobes, each at least 1 mm wide (Beetle 1959, 1960). Flowerheads bear three to 11 disc flowers and are arranged into leafy, narrowly General Rabbitbrush Culture ______racemose panicles. Flowers bloom in late August and Table 4 presents distribution and differentation of September. Seed ripens in October. Wyoming threetip rabbitbrush taxa. Tueller and Payne (1987) and other sagebrush has a rather limited range. It occurs on papers in Johnson (1987) provide additional informa- rocky knolls from 7,000 to 9,000 ft (2,100 to 2,700 m) tion on rabbitbrush culture, management, and use. in elevation in central and southeastern Wyoming The fruit of rabbitbrush is a spindle shaped achene (Beetle 1960). Brunner (1972) reported this subspe- (seed). The terminal, or crown end, of each seed bears cies also occurs in southern Oregon, but had not yet a ring or crown of hairs, known as the pappus. Seed can been found in Nevada. It typically grows on sites be wind and animal dispersed. Normally seed is pro- adjacent to those of mountain big sagebrush. duced annually. Seedlings naturally establish, espe- Tall threetip sagebrush (A. t. ssp. tripartita) is a cially on disturbed sites where competition is lacking freely branching shrub up to 6 ft (1.8 m) high. It can or slight. Thousands of seedlings can be found in bare layer easily when the conditions are right but is and disturbed soils, under and next to mature plants. seldom found layering in the field. After burning, it Most do not, however, develop beyond the seedling may stump-sprout (Beetle 1960). Leaves of the vegeta- stage. Seeds are relatively small (see chapter 20) with tive branches are 1.5 to 4 cm long and deeply divided considerable variation in size between taxa. White into three linear lobes less than 1 mm wide. The lobes rubber rabbitbrush seeds are among the smallest with may be further divided (Beetle 1959, 1960). about 700,000 per lb (1.5 million/kg) (100 percent Flowerheads bear four to eight disc flowers and are purity). Mountain rubber rabbitbrush seeds are some- arranged into panicles that may sometimes be re- what larger with just over 425,000 per lb (937,000 per kg). duced to a spicate form. Flowers bloom in late August Various rabbitbrush species have been successfully and September. Seeds ripen in October. This subspe- seeded. Seeding success, based on successful estab- cies occurs in dry, well drained soils at 3,000 to 7,500 ft lishment, has ranged from 0.001 to 4.0 percent (Stevens (910 to 2,300 m) elevation from British Columbia and others 1986). The pappus has to be removed to south through Washington to northern Nevada and facilitate seed being seeded aerially, broadcast, or eastward to northern Utah and western Montana. drilled. Although seeding success theoretically could be

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Table 4—Species of Chrysothamnus with distribution, notes, and subspecific differentiation (drawn from Anderson 1986a; McArthur and Meyer 1987; McArthur and others 1979a; and references cited therein).

Section and Number of species Distribution Notes subspecies Chrysothamnus C. albidus Scattered in alkaline soils W Utah to White-flowered very resinous — EC California.

C. greeneia Often in sandy soils SC Colorado and N Mature plants have brittle white shiny stems 1 New Mexico to W Nevada and E California. Involuvral bracts are acuminate and recurved.

C. humilis Arid plains NW Nevada, NE California, Blooms earlier than rabbitbrush. — SE Oregon, and isolated populations further N in Oregon and Washington

C. linifolius Dry arroyos and other moist, deep but Large statured like rubber rabbitbrush. — usually alkaline soil sites S Montana to N Arizona and N New Mexico

C. viscidiflorus Broad distribution on dry open areas British Some populations have the only ray 5b Columbia and Montana S to New Mexico, flowers in Chrysothamnus Arizona, and E California; entire state of Utah. Gramini C. eremobius Known from only three populations in upland Related to genus Petradoria — S Nevada.

C. gramineus Grows in mountain canyons in S Nevada Related to genus Petradoria. As opposed — and adjacent California. to C. eremobius this species has longer involucres, unkeeled phyllaries, and glabrous achenes. Nauseosi C. nauseosus Broadly distributed in plains, foothills and Most common and widely distributed 22c mountains British Columbia to Saskatchewan rabbitbrush with much diversity. Forms of S to Western Texas, Sonora, and Baja ssp. albicaulis, hololeucus, and salicifolius California. are preferred by browsing animals.

C. parryi Dry open places in mountains and foothills Shares stem tomentum character with Wyoming and Western Nebraska W to C. nauseosus but has attenuate involucral 13d California and S to New Mexico and Arizona. bracts as opposed to acute bracts of C. nauseosus. Generally smaller and finer structured than C. nauseosus. Pulchelli C. depressus Dry plains, hills, and rocky mountain slopes Dwarf shrub with conspicuously keeled, — scattered over Western Colorado and New sharply ranked involucral bracts. Perhaps Mexico W to SE California and S Nevada. the most palatable taxa to ruminants.

C. molestus Occurs only in a few scattered populations Rare species. Has glandular-hispid foliage. — in uplands of NC Arizona.

C. pulchellus An undershrub in SW desert savannas SE Utah Has prominantly angled achenes. — and SW in Utah, Kansas, to W Texas and N Coahuila.

C. vaseyi Scattered populations in mountain valleys and Resembles the more common C. viscidiflorus.— foothills SE Wyoming to NC New Mexico W to high plateaus of Utah.

Punctati C. paniculatus Arid often stony slopes especially in dry arroyos Striate stems and terete leaves. — SW Utah and SW Nevada to SC California.

C. teretifolius Arid stony slopes in S Nevada and SE California. Similar to C. paniculatis but more abundant. — Has more strongly aligned involucral bracts

aFormerly ssp. greenei and filifolius were recognized, but McArthur and others (1978a, 1979a) and Anderson (1986a) see no consistent distinctive differences between the formerly recognized subspecies. b A total of 25 have been named at various taxonomic ranks. Hall and Clements (1923) recognized nine; Anderson (1986a) recognized five subspecies: ssp. axillaris, lanceolatus, planifolius, puberulus, and viscidiflorus. c A total of 62 have been named at various taxonomic ranks. Hall and Clements (1923) recognized 20; Anderson (1986a) recognized 22 subspecies: ssp. albicaulis, arenarius, bernardinus, bigelovii, ceruminosos, graveolens, hololeucus, iridis, junceus, latisquameus, leiospermus, mohavensis, nanus, nauseosus, nitidus, psilocarpus, salicifolius, texensis, turbinatus, uintahensis, and wasinoensis. dA total of 16 have been named at various taxonomic ranks. Hall and Clements (1923) recognized 10; Anderson (1986a) recognized 12 subspecies: ssp. affinis, attenuatus, asper, howardii, imulus, latior, monocephalus, montanum, nevadensis, parryi, salmonensis, and vulcanicus.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 523 Chapter 21 Composite Shrubs enhanced where the pappus is not removed (Stevens and others 1986), there is presently no available equipment for seeding uncleaned seed. Seeds are generally hand collected but can be harvested with various types of seed strippers, seed beaters, and vacuums. The pappus is removed by running the seed through a barley debearder or hammermill followed by screening and fanning. Seed is generally collected between 10 and 15 percent purity. Seed can be cleaned to 95 percent purity with persistent screening and fanning. Barley debearding is preferred over hammermilling. Seeds are somewhat brittle; hammermilling can result in seed breakage and crack- ing in the balling of the collected material. Balling hinders seeding. Seed cleaned to less than 70 percent purity can only Figure 34—Dwarf rabbitbrush growing near the be seeded aerially, by hand, or through a thimble mouth of Ephraim Canyon, Sanpete County, UT. seeder. Seed cleaned to 70 percent or greater can be run through a drill, seed dribbler, or browse seeder. Where seeds are cleaned to high purities, they need to be diluted with rice hulls or screened sawdust to facili- This species occurs on dry plains, hills, and rocky tate seeding. Seed of most rabbitbrush species should mountain slopes from 3,300 to 6,900 ft (1,000 to 2,100 be surface seeded or covered not more than 1⁄32 inch (0.8 mm) deep. If drilled, seed drop tubes and hoses m) in elevations scattered over western Colorado, New should be removed from between the disk furrow Mexico, Utah, Nevada, Arizona, and southeastern opener and placed behind them. Seed will then be California (Anderson 1986a). deposited behind the disk furrow opener on a dis- Dwarf rabbitbrush is often heavily browsed by sheep, turbed soil surface. Rabbitbrush seedlings are gener- cattle, and wildlife (Hall and Clements 1923; Kufeld ally not competitive and may be suppressed by other and others 1973). It transplants readily, can be seeded, species. If drilled, thimble, dribbler, or browse seeder and is useful for stabilizing depleted soils on which it seeded, rabbitbrush seed should be seeded individu- readily grows. This handsome shrub is a source of ally or with less competitive species. Broadcasting protein when grasses and broadleaf herbs have dried results in substantially less competition between spe- (McArthur and others 1979a). cies. Late fall and early spring seeding are preferred. Rabbitbrush seed can be stored in an open, unheated, uncooled warehouse for up to 3 years following collec- Chrysothamnus linifolius ______tion without too much loss of viability. Spreading Rabbitbrush Chrysothamnus depressus ______Spreading rabbitbrush is a tall, robust shrub up to 7 ft (2.1 m) high that spreads underground (fig. 35) by Dwarf Rabbitbrush lateral roots that form adventitious shoots (Anderson 1964). Of all the rabbitbrushes only Parry rabbitbrush Dwarf rabbitbrush is a small, irregularly branched, shares the underground spreading trait and then not depressed shrub or subshrub 1 ft (30.5 cm) high or less, nearly to the extent spreading rabbitbrush does. Leaves with numerous short branches arising from decum- are large, flat, green, glabrous, lanceolate to oblong- bent lower stems (fig. 34). The branches are covered lanceolate, 2 to 5 cm long, and 8 to 16 mm wide. Heads with a dense scabrid pubescence. The narrowly oblan- contain four to six, usually five, disc flowers and are ceolate to spatulate leaves are 0.7 to 2 cm long, 1 to arranged into broad, loose cymes. The involucral bracts 4 mm wide, and finely puberulent or scabrous like the have obtuse tips with thickened green spots similar to branches. The heads contain five disc flowers that those of Vasey rabbitbrush. These spots, however, dry are arranged into compact terminal cymes. Involucral to a brown color. Achenes are covered with dense, long, bracts have mucromate to attenuate tips, are 10 to soft hair. Blooming occurs during August and Septem- 13 mm long, and are arranged into five distinct ber with seed ripening from October to mid-December. vertical ranks. Achenes are 5 to 5.5 mm long and Spreading rabbitbrush occurs in Montana, Wyoming, glabrous or obscurely pubescent toward their apex. Colorado, New Mexico, Utah, and Arizona. It is most The soft brownish-white pappus is slightly longer than abundant in alkaline soils along roadcuts, barrow pits, the corolla. Blooming occurs from May to October; seed ditches, streams, and washes in the Upper Colorado matures from October to December (see chapter 20). River drainage.

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Figure 35—Root sprouts, foreground, of spread- Figure 36—Large rubber rabbitbrush growing at ing rabbitbrush at the Snow Field Station. Palmetto, Esmeralda County, NV. The specimen pictured is the viridulus form of ssp. consimilis.

The species has aggressive underground spreading the end of the year with over 500,000 cleaned seeds characteristics coupled with an ability to spread by per lb (1.1 million/kg) (see chapter 24). Rubber rabbit- seed that make it a valuable stabilizer of disturbed brush is normally a heavy seed producer (fig. 37). alkaline soils (McArthur and others 1974; 1979a; Plummer 1977). It transplants readily. Laycock and Shoop (1986) found spreading rabbitbrush to be an Ecological Relationships and excellent living snowfence for the central Great Plains. Distribution ______Spreading rabbitbrush receives little browsing use. It does, however, provide year round cover for upland We have recently discovered that galls induced by game and small birds. tephritid flies are differentially distributed among the subspecies of rubber rabbitbrush (McArthur 1986; McArthur and others 1979b—See Varieties and Chrysothamnus nauseosus ______Ecotypes section later in this section for subspecies descriptions). Over much of their range, the white- Rubber Rabbitbrush stemmed subspecies albicaulis and subspecies Rubber rabbitbrush is a shrub usually 12 to 80 hololeucus are infested with a persistent round stem inches (0.3 to 2.0 m) high, but it varies from dwarf gall, 0.3 to 1.2 cm in diameter. This gall was found forms to types over 10 ft (3.0 m) high (fig. 36). Usually, restricted to these white-stemmed subspecies in our several erect stems arise from the base and these observations. Green subspecies consimilis, graveolens, branch to form rounded bushes. Branches are covered with a green, yellow-green, gray-green to white feltlike tomentum usually infiltrated with a resinous gum, making the plant somewhat sticky. This coating is often mistaken for part of the bark but can be discerned by scraping with a knife edge or a fingernail. Leaves are nearly filiform in some subspecies to broadly linear in others. Leaves vary from 18 to 63 mm long and are covered with a tomentose vestiture. They are not twisted or gland-dotted. The heads of this species are usually arranged into a cymose inflorescence. Each head bears 20 to 25 glabrous to densely tomentose involucral bracts arranged in up to five vertical rows. Rubber rabbitbrush blooms from August to October and are among the latest bloomers of the genus. Plants at higher elevations bloom earlier than those at lower elevations. Seeds mature, depending on location and Figure 37—Rubber rabbitbrush in seed, south- central Utah. subspecies, and are collected from mid-October until

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 525 Chapter 21 Composite Shrubs and turbinatus, on the other hand, have a less persistent, fluffy stem gall reminiscent of a ball of cotton about 0.7 to 1.4 cm in diameter. This gall is found also on ssp. hololeucus in southern areas near Kanab, UT, and Colorado City, AZ. The round gall is absent in these areas. In areas where these green-stemmed and white-stemmed rubber rabbitbrush subspecies occur together the galls are specific. We have observed only a few cases of cross gall inoculation and then only on putative hybrid plants. Wangberg (1981) independently observed some species and subspecies specificity of tephritid galls on rabbitbrush in Idaho. He found less specificity for the round gall in Idaho than we did in Utah. He identified the tephritid flies that induce the galls as two different species of Aciurina. Dodson and George (1986) confirmed the genetic and ecological integrity of the two Aciurina species. Rubber rabbitbrush ranges from British Columbia Figure 38—Rubber rabbitbrush growing in as- to Saskatchewan south to western Texas, Sonora, sociation with smooth brome, Sanpete County, UT. Note vigor of grass under shrub canopy. Baja California, and eastern California (Anderson 1986a, c). It is a common plant on plains, valleys, and foothills. It grows best in openings with the sagebrush, juniper-pinyon, and ponderosa pine zones in sandy, gravelly, or clay-alkaline soils. This species grows or early winter. Once established, natural spread can at elevations ranging from 500 to 9,000 ft (150 to occur (Stevens 1986b). 2,700 m). Rubber rabbitbrush is a vigorous early Seed germination of rubber rabbitbrush is usually invader of disturbed sites such as roadcuts and over- quite high. Seeds maintain high viability (65 percent) grazed rangelands. On ranges where big sagebrush for 3 years under ordinary warehouse storage condi- has been destroyed by fire, insects, vehicular traffic, or tions (Stevens and others 1981a). With cool night continued heavy grazing, rabbitbrush will increase (40.5 ∞F; 4.7 ∞C) and warm day temperatures, rubber and often become the dominant vegetation (Evans and rabbitbrush seed germinates in about 2 days (Weber others 1973; Rosentreter 1986). Nevertheless, in most and others 1985). However, under standardized single habitats, this species is not overly competitive with cool temperature germination conditions, seeds from herbaceous species, and on some sites, it does not lower elevations and latitudes germinate in about suppress grass. Production of herbaceous cover per- half the time (2 weeks) that it takes for seeds from centage have been notably greater (fig. 38) when higher elevations and more northerly areas to germi- rabbitbrush is present than when it is not present nate (McArthur and others 1987a; Meyer and McArthur (Frischknecht 1963; Plummer 1959; Plummer and 1987; Meyer and others 1989). Stevens and others others 1968). We find it of interest that rubber rab- (1986) have shown that seeding rabbitbrush is more bitbrush has a high photosynthetic rate, not unlike successful when the pappus is left on the achene (the desert-adapted C4 photosynthetic pathway species fruit that holds rabbitbrush seed) and the achenes are (Davis and others 1985). placed upright at the soil surface. Achene size in rubber rabbitbrush subspecies appears to be corre- Plant Culture lated to habitat; subspecies adapted to sandy sites have larger achenes (Meyer 1997). Rubber rabbitbrush can easily be transplanted in the fall or preferably in the spring to avoid frost Uses and Management heaving so long as the plants are physiologically inac- tive and moisture conditions are right and the trans- Rubber rabbitbrush is an excellent plant for con- plants are small (10 inches tall; 25.4 cm or less). A less trolling erosion because of its deep roots, heavy litter, successful method—which, nevertheless, yields some and ability to establish on severe sites (Aldon and Pase results—is to divide dormant crowns (Hall and 1981; Luke and Monsen 1984; Monsen and Richardson Clements 1923). Tissue culture techniques have been 1984; Monsen and Stevens 1987; USDA Forest Service successfully employed to propagate rubber rabbit- 1937). Once established, this species reproduces brush (Upadhyaya and others 1985). Success was easily and spreads fast from its light, plumed, wind- obtained from direct seeding, especially when seeds disseminated achenes (Young and others 1984b). It have been distributed on top of disturbed soil in the fall also grows vigorously when transplanted. The value of

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C. nauseosus as browse varies greatly between sub- A species and ecotypes. The white to grayish subspecies such as albicaulis, hololeucus, and salicifolius are more palatable to livestock and big game than the green subspecies, graveolens and consimilis (fig. 39) (Hanks and others 1975; McArthur and others 1974). Throughout much of the summer range, game and livestock browse the plant lightly, if at all, except under unusual conditions. During late summer and fall when rubber rabbitbrush is in bloom, most livestock and game graze the flowers and occasionally a few leaves and the more tender stems. Rubber rabbitbrush is most heavily browsed on winter ranges (fig. 40). Rubber rabbitbrush was found in 48 percent of the deer stomachs examined on a portion of winter range in Owens Valley, Inyo County, CA; however, it never amounted to more than 6 percent of the total food ingested (Sampson and Jesperson 1963). Crude protein content ranged from 9 percent during the B dormant months to 11.8 percent in the spring after new leaves had formed (Sampson and Jesperson 1963). Antelope also make extensive use of rubber rabbitbrush (Yoakum 1986). Bhat and others (1990) found a wide range of winter nutritive quality in rubber rabbitbrush accessions growing in a uniform garden. Rubber rabbitbrush is resilient to browsing. Severely browsed plants often obtain full regrowth by midsummer following heavy browsing. Rubber rabbitbrush also provides cover for small mammals and birds. The potential for this species to produce rubber and other chemicals has long been recognized (Hall and

Figure 40—Heavy use of rubber rabbitbrush by deer in the winter at an experimental plot in Sanpete County, UT: (A) C. nauseosus spp. salifolius plant in the late fall; (B) The same plant after winter browsing. The stake in “A” is 1 ft tall; the mark on the stake in “B” is at 1 ft.

Goodspeed 1919). There has been a recent revival of interest in this potential (Hegerhorst and others 1987; Ostler and others 1986; Weber and others 1985). Some accessions produce up to 6 percent stem rubber and 20 percent resin content. The plant also has other possible industrial and horticultural uses (Weber and others 1985). Rubber rabbitbrush can become troublesome on ranges when it invades and occupies disturbed lands Figure 39—Differential winter-spring utiliza- at high densities. It is a difficult plant to control tion of rubber rabbitbrush subspecies by deer in Sanpete County, UT. Left, C. nauseosus because of its resistance to herbicides and its crown ssp. consimilis (10 percent use), right, C. sprouting nature (Evans and others 1973; Whisenant nauseosus ssp. hololeucus (50 percent use). 1986b, 1987; Young and others 1976a).

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Varieties and Ecotypes Anderson (1986a,c) recognized some 22 subspecies of rubber rabbitbrush. Often, two or more subspecies will occur sympatrically. Each subspecies is essentially inbred and morphologically clean (Anderson 1986c; McArthur and Meyer 1987; McArthur and others 1978a). However, reproductive isolation occasionally breaks down, especially at the limits of subspecies geographical ranges, and intermediate morphological forms are found (Anderson 1986c; Hanks and others 1975; McArthur and others 1978a). Be- cause of the mostly inbreeding, but potential for out- breeding, system of rubber rabbitbrush, many local C reproductively isolated populations of rubber rabbitbrush occur. We agree with Hall and Clements (1923) that “improvement in any desired direction may be brought about by selection, or by hybridization or by both of these methods.” To date, no artificial improvement has been made, but individual populations A with desirable characteristics such as high rubber content, palatability, or growth form have been iden- D tified. We discuss below the more common subspecies B of rubber rabbitbrush. White stem rubber rabbitbrush—C. nauseosus ssp. Figure 41—Four subspecies of rubber rabbit- albicaulis (mountain), and hololeucus (basin)—are brush: (A) C. nauseosus ssp. albicaulis; (B) C. shrubs from 2 to 6.5 ft (0.6 to 2.0 m) high with erect, nauseosus consimilis; (C) C. nauseosus ssp. leafy branches and leaves covered with a permanent, graveolens; (D) C. nauseosus ssp. salicifolius dense, white to grayish tomentum (fig. 41). The leaves (McArthur and others 1979a). are 2.5 to 4 cm long, 0.5 to 1.5 mm wide or sometimes to 3 mm wide in a few forms. The strongly keeled, acute involucral bracts are white, more or less tomentose, hololeucus is often intermixed with ssp. graveolens 8 to 10 mm long, and arranged into five distinct on foothill ranges and with subspecies consimilis in vertical ranks. The yellow corollas are 8 to 11 m long valleys and plains. The striking white forms of both and terminate in lobes 1 to 2 mm long. The achenes are subspecies have potential use as ornamentals densely pubescent. The subspecies differ in the length (McArthur and others 1979a; Weber and others 1985). of their corolla lobes (ssp. albicaulis 1 to 2 mm long, Another common white subspecies of the foothills and ssp. hololeucus 0.5 to 1 mm long) and style appendage plains of Colorado, Wyoming, Montana, and the west- lengths for which ssp. albicaulis has style appendages ern Dakoatas is ssp. nauseosus. This subspecies is longer than stigmatic portions, but ssp. hololeucus short statured, about 2 ft (61.0 cm) tall, and quite does not (Anderson 1986a). These subspecies tend to palatable to game and livestock. overlap morphologically in their area of sympatry Threadleaf rubber rabbitbrush (C. n. ssp. consimilis) (northern Utah, northeastern and western Nevada, may reach 10 ft (3.0 m) in height when mature. It has southwestern Idaho, and southeastern California; leafy, erect branches covered with a green to yellow- Anderson 1986a,c). In these areas of sympatry, sub- green tomentum. The narrow threadlike (linear- species albicaulis occurs at higher elevations (over filiform) leaves are less than 1 mm wide and 2.5 to 5 cm 6,000 ft; 1,800 m) whereas ssp. hololeucus is confined long (fig. 41). They are usually covered with a green to to lower elevations. White rubber rabbitbrush subspe- yellow-green tomentum and are somewhat resinous. cies are common and widespread and are found in The involucral bracts are acute, glabrous, and keeled open places in plains, foothills, and mountains from and are arranged in fairly distinct vertical rows. The British Columbia, Alberta, and Montana southward bracts are 6.5 to 8.5 mm long. The corollas are 7 to 9.5 to northwestern Colorado, Utah, Nevada, and southern mm long, with glabrous lobes 1 to 2.5 mm long. The California (Anderson 1986c). Subspecies hololeucus achenes are densely pubescent, suggesting that this grows in the Great Basin area whereas ssp. albicaulis subspecies may be a connecting link between rubber occurs on mountains and foothills, east, north, and rabbitbrush and low rabbitbrush. Subspecies con- west fringes of the Great Basin and beyond (Anderson similis is most common in alkaline valleys and plains 1986a,c; Hall and Clements 1923). Subspecies of the Great Basin where it is often associated with

528 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs various saltbushes and black greasewood. In less widespread at higher elevations in Utah and perhaps alkaline areas, consimilis intermixes with ssp. extending to Nevada as part of the lower subalpine hololeucus and graveolens. Threadleaf rubber rabbit- vegetation. It extends down to the foothills in parts brush also occurs in alkaline soil outside the Great of its range where it may be found intermixed with Basin from western Wyoming, Colorado, and New ssp. albicaulis, hololeucus, and graveolens (Plummer Mexico to northeastern Oregon and eastern California. 1977). Mountain rubber rabbitbrush appears to be A large ecotype, up to 12 ft (3.7 m) tall with treelike the most palatable subspecies for both livestock and trunks, occurring principally in the western parts of big game of the five common subspecies, salicifolius, its range, was formerly recognized as a separate sub- albicaulis, hololeucus, graveolens, and consimilis in species (viridulus) (fig. 36). Perhaps ssp. viridulus Utah, and it is often browsed heavily in the fall should be maintained as it is higher in rubber content (McArthur and others 1979a). and in resins than ssp. consimilis (Ostler and others Other less common subspecies with locally large 1986). Another subspecies, pinifolius, has also been populations include arenarius, junceus, leiospermus, reduced to be synonymous with consimilis (Anderson and turbinatus. Subspecies arenarius is a striking 1986a). Plants formerly referred to as pinifolius grow white shrub adapted to sand dunes in the Four Cor- in southern Colorado and New Mexico. Threadleaf ners area. It has large heads up to 20 mm long. Sub- rubber rabbitbrush is one of the least palatable of the species junceus is a nearly leafless, yellow-green form C. nauseosus subspecies. It may, therefore, have value adapted to sandy areas in the Colorado River drain- in revegetating disturbed sites such as roadcuts where age. Subspecies leiospermus is a low shrub usually attraction of browsing animals is not desired. This less than a meter in height. It has affinities to ssp. subspecies helps control erosion on open alkaline soils consimilis. It occurs mostly on arid, rocky sites in the by providing ground cover and soil stabilization southern half of the Intermountain area. A distin- (McArthur and others 1979a). guishing feature of ssp. leiospermus is its glabrous or Green rubber rabbitbrush (C. n. ssp. graveolens) nearly glabrous achenes. Subspecies turbinatus is a ranges from 2 to 5 ft (0.6 to 1.5 m) high when mature. yellow form adapted to sandy sites, mostly in western Its leafy, erect branches are yellow-green to green or Utah. It has large floral heads. A few additional sometimes gray-green and are covered with a compact subspecies occur principally outside of the Intermoun- tomentum (fig. 41). The linear leaves are 1 to 3 mm tain area. These include bigelovii (mostly south and wide, 4 to 6 cm long, and only slightly pubescent. The west of the Four Corners area), nauseosus (mostly on involucral bracts are 6 to 8 mm long, glabrous at least the western edge of the northern Great Plains), and on their backs, acute, keeled, and arranged in vertical mohavensis (southern California) (Anderson 1986a,c; rows. Achenes are densely pubescent. Green rubber McArthur and Meyer 1987; McArthur and others rabbitbrush is widespread and sporadic from North 1979a). Dakota to Idaho and southward to western Texas, New Mexico, and Arizona. It is most common on well- drained foothills but also extends up into the moun- Chrysothamnus parryi ______tains and down into valleys and plains where it is Parry Rabbitbrush often intermixed with ssp. consimilis. Green rubber rabbitbrush is generally less palatable than the white Parry rabbitbrush (Chrysothamnus parryi) is a or gray subspecies, hololeucus, albicaulis, and shrubby species somewhat intermediate in height, salicifolius. Nevertheless, some forms of this subspe- stem and leaf tomentum, and growth habit between cies have been found that are utilized to a moderate rubber rabbitbrush and low rabbitbrush. Parry rab- degree by livestock and mule deer. bitbrush is a low, dense shrub similar in habit to Mountain rubber rabbitbrush (C. n. ssp. salicifolius) certain forms of low rabbitbrush (fig. 42). It is usually is a shrub from 1 to 6.5 ft (0.3 to 2.0 m) high. Its from 8 to 24 inches (20.3 to 61.0 cm) in height with ascending to erect twigs are leafy and covered with a numerous spreading to erect flexible branches. Like gray-green, fairly compact tomentum. The leaves are rubber rabbitbrush, the branches of Parry rabbit- broadly linear, ranging from 4 to 8 cm long and 3 to brush are covered with a feltlike white to green tomen- 10 mm wide, which makes them the largest leaves of tum. The tomentum is neither as dense nor resinous the species (fig. 41). The involucral bracts are 7 to 8 mm as in rubber rabbitbrush. Paulsen and Miller (1968) long, mostly obtuse, nearly glabrous, and arranged and McArthur and others (1979a) observed that Parry in rather obscure ranks. The yellow corollas are about rabbitbrush spread from underground roots. The 1 cm long and have a minutely pubescent throat. The glabrous to tomentose, somewhat viscid leaves are achenes are densely pubescent. Hall and Clements narrowly linear to elliptic and range in size from 0.5 to (1923) reported that mountain rubber rabbitbrush is 8 mm wide and 1 to 8 cm long. Flowerheads usually are “apparently rare and confined to Utah.” However, arranged in terminal leafy racemes that sometimes McArthur and others (1979a) found it to be fairly form panicles. The involucral bracts are 9 to 14 mm

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Chrysothamnus parryi ssp. attenuatus consists of low shrubs with mostly erect stems up to 2 ft (61.0 cm) high. It has green, slightly viscid, narrowly linear leaves, 2 to 4 mm long and about 1 mm wide. The leaves are erect but are not larger than the inflorescence. Heads contain five to seven disc flowers and are subtended by involucral bracts with slender, straight tips. The bracts are ranked into five vertical rows. Blooming occurs from August to October. Sub- species attenuatus is found in the sagebrush, pinyon- juniper, and yellow pine vegetation types in Utah and southwestern Colorado and northwestern New Mexico (Anderson 1986a; McArthur and others 1979a). Chrysothamnus parryi ssp. howardii is a low shrub. Its spreading basal stems and erect branches Figure 42—Parry rabbitbrush (C. parryi ssp. are up to 2 ft (61.0 cm) high. The narrowly linear, howardi) growing at Current Creek, Duchesne tomentose leaves are 2 to 4 cm long, about 1 mm wide, County, UT. and the upper ones usually extend beyond the uppermost heads of the inflorescence. Flowerheads contain five to seven pale yellow disc flowers. The heads are subtended by vertically ranked involucral bracts usually with spreading tips. Blooming occurs from high and terminate in acuminate to very attenuate July to September. The subspecies occurs on dry hills herbaceous tips. The yellow, tubular to funnelform and mesas associated with sagebrush, pinyon-juniper, corollas are 8 to 11 mm long. Achenes are 5 to 6 mm and ponderosa pine vegetational types in Utah, south- long and are covered with long, shaggy, appressed ern Wyoming, Colorado, New Mexico, and Nebraska hairs. Blooming occurs from July to September; seed (Anderson 1986a; McArthur and others 1979a). matures in October and November. Chrysothamnus parryi ssp. monocephalus is a low shrub from 4 to 24 inches (10.2 to 61.0 cm) high with Ecological Relationships and Distribution rigid, spreading branches. Its viscid, somewhat Parry rabbitbrush occurs in dry, open places in tomentose leaves are linear-oblanceolate or spatulate, mountains and foothills of Western North America 1 to 3 cm long and 1.5 cm or less wide. The upper leaves from Wyoming and western Nebraska west to usually extend beyond the inflorescence. The flower- California and south to New Mexico and Arizona. ing heads occur singly or in pairs on the end of short, Like other species of rabbitbrush, this species tends leafy branches. The heads contain five to six disc to increase on overgrazed and disturbed areas. flowers and are subtended by obscurely ranked in- Populations of Parry rabbitbrush are usually smaller volucral bracts with straight, attenuate tips. This and more scattered than those of the other common subspecies occurs in the high mountains of western species (rubber and low rabbitbrushes) of rabbitbrush. Nevada and eastern California at elevations between 2,600 and 11,000 ft (790 and 3,400 m) (McArthur and Varieties and Ecotypes others 1979a). Nevada rabbitbrush (Chrysothamnus parryi ssp. Parry rabbitbrush is a diverse widespread group, nevadensis) consists of low shrubs with ascending with 12 subspecies (Anderson 1986a; McArthur and to erect branches up to 2 ft (61.0 cm) high. The linear Meyer 1987). to linear-oblanceolate leaves are 1.5 to 4 cm long, Chrysothamnus parryi ssp. asper is a low shrub 0.5 to 3 mm wide, and sometimes green to resinous, 6 inches (15.2 cm) or more high with slightly spread- but usually gray-tomentose. The uppermost leaves ing to erect branches. Its green leaves, roughened rarely extend beyond the inflorescence. The flowering with short-stalked resin glands, are 2 to 5 cm long and heads contain four to six yellow disc flowers and are 1 to 3 mm wide. The heads contain five to 10 disc subtended by ranked involucral bracts with slender flowers and are subtended by somewhat ranked in- recurved tips. Chrysothamnus parryi ssp. nevadensis volucral bracts with straight tips. This subspecies occurs between 4,300 and 8,900 ft (1,300 and 2,700 m) occurs on mountain sides bordering desert areas in elevation on dry mountain sides from eastern from 6,900 to 8,500 ft (2,100 to 2,600 m) in elevation in California to eastern Nevada, southwestern Utah, western Nevada and eastern California (Anderson and northern Arizona. It is most common along the 1986a; McArthur and others 1979a). eastern flank of the Sierra Nevada (Anderson 1986a; McArthur and others 1979a).

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Chrysothamnus parryi ssp. parryi consists of low and often twisted. Leaf vestiture is glabrous or pu- shrubs with erect branches 1 to 2.5 ft (30.5 to 76.2 cm) bescent and commonly viscidulous with usually high. The uppermost leaves usually extend beyond scabrous margins. Degree of pubescence may vary the inflorescence. Flowering heads contain 10 to 20 disc tremendously in variants of this species. As Hall and flowers and are subtended by obscurely ranked in- Clements (1923) pointed out, sometimes the pubes- volucral bracts with straight attenuate tips. It blooms cence among the subspecies of low rabbitbrush “will during August and September. Subspecies parryi grows occur as a fairly dense though minute puberlence in in dry plains, valleys, and hillsides in central Nevada, certain plants, while others almost exactly dupli- southern Utah, south-central Wyoming, western Colo- cating these in every other respect will be perfectly rado, and northern New Mexico (Anderson 1986a; glabrous.” Furthermore, L. C. Anderson (n.d.) has McArthur and others 1979a). observed good correlation between plant stature and A few other subspecies of C. parryi, such as affinis, leaf size of ssp. visicidiflorus and lanceolatus with glandulosa, imulus, and vulcanicus, occur outside of altitude and amount of precipitation. Flowerheads or only on the fringe of the Intermountain area and in small numbers (Anderson 1986a; Hall and Clements containing approximately five perfect, fertile disc 1923). No efforts have been made to improve Parry flowers each are arranged in compact terminal cymes. rabbitbrush. The species contains abundant genetic Involucral bracts number about 15 per head and are variation. Its ssp. howardii is thought to be involved arranged in poorly to well defined vertical ranks. The in the parentage of C. nauseosus ssp. utahensis bracts of some subspecies have a greenish or brownish (Anderson 1984). thickened spot near their apex (McArthur and others 1979a). Flowering occurs from August through Octo- ber; seed matures from October to the end of Decem- Chrysothamnus vaseyi ______ber. Low rabbitbrush produces about 3.4 million seeds Vasey Rabbitbrush per lb (7.5 million/kg) of cleaned seed (see chapter 24). This species has strong basal sprouting tendencies, Vasey rabbitbrush is a low, rounded shrub with especially following top removal and injuries (Wasser ascending to erect branches up to 1 ft (30.5 cm) high. 1982). The bark on the young branches is pale green to whitish and glabrous, becoming brown and fibrous Ecological Relationships and Distribution with age. Leaves are linear to linear-oblanceolate, 1 to 2.5 cm long, 1 to 3 mm wide, and glabrous. Heads contain Low rabbitbrush is one of the most widely distrib- five to seven disc flowers each and are arranged into uted shrubs on rangeland throughout Western North small, compact cymes. The obscurely ranked involu- America (Anderson 1986a,b; McArthur and Meyer cral bracts are 5 to 7 mm high, oblong, obtuse to 1987). The species has great ecological amplitude. It rounded, and all but the innermost have a thickened occurs in dry, open areas from British Columbia and greenish spot near the apex. The achenes are about Montana, south to New Mexico, Arizona, and eastern 5 mm long, terete, longitudinally 10-striate, and glabrous. Blooming occurs from July to September; seed matures from October until mid-December (table 1). Vasey rabbitbrush occurs scattered over plains, hillsides, and mountain valleys at altitudes of 5,600 to 8,500 ft (1,700 to 2,600 m) mostly in Utah and Colorado, but also in New Mexico and Wyoming (Anderson 1986a; Hall and Clements 1923). This shrub is browsed by sheep but is small and so scattered that it provides little forage (McArthur and others 1979a).

Chrysothamnus viscidiflorus______Low Rabbitbrush Low rabbitbrush varies in appearance and foliage characteristics. It is usually 1 to 3.5 ft (0.3 to 1.1 m) tall with many erect stems branching from a simple Figure 43—Low rabbitbrush growing at the Snow base (fig. 43). The brittle, erect twigs are glabrous or Field Station, Sanpete County, UT. This speci- puberulent with pale green or white bark. Leaves are men is a mountain low rabbitbrush (C. viscidiflorus narrowly linear to oblong or lanceolate, 1 to 6 cm long ssp. lanceolatus).

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California at elevations between 2,600 and 11,000 ft Varieties and Ecotypes (790 and 3,400 m). Low rabbitbrush is usually associated with sagebrush, snakeweed, and other species of This species includes five subspecies and several rabbitbrush. Anderson (1986b) has shown that poly- ecotypes within subspecies (Anderson 1980, 1986b; ploid races in those subspecies where polyploidy oc- McArthur and Meyer 1987). Most important among curs (ssp. lanceolatus, puberulus, viscidiflorus) are these in the Great Basin are two glabrous subspecies, viscidiflorus and axillaris, and two pubescent sub- adapted to lower and drier sites than their diploid species, lanceolatus and puberulus. Anderson (1980) counterparts (Anderson 1986b). reduced ssp. stenophyllus to varietal status under ssp. viscidiflorus. Hall and Clements (1923) believe nu- Plant Culture merous intergrades have been held together in one Low rabbitbrush can be transplanted in the fall rather close species through interbreeding where their ranges meet or overlap. Abrams and Ferris and spring. When transplanting in the fall consider- (1960) describe C. viscidiflorus as a highly polymorphic ation needs to be given to frost heaving potential. species composed of several freely intergrading sub- Fall seeding is preferred; seed can be drilled or broad- 1 species of overlapping distribution. Intermediates are cast, and seed should not be covered more than /8 inch invariably found in the field. We believe forms of low (3.2 mm). Seeds can be harvested by hand and by rabbitbrush may be improved for grazing and other vacuum harvesting techniques. uses through selection and breeding. Because each subspecies is self-fertilized and predominately self- Uses and Management pollinated, each maintains its identify despite occasional outcrossing (McArthur and others 1978a). This shrub may provide an important supply of Mountain low rabbitbrush (Chrysothamnus viscid- browse to both game and livestock, particularly dur- iflorus ssp. lanceolatus) is a small shrub from 8 to ing late fall and winter after more desirable forage 20 inches (20.3 to 50.8 cm) tall (fig. 43, 44). Its branches has been consumed. Throughout the Great Basin, low are straw colored or gray and are finely pubescent. rabbitbrush, especially the flowering shoots, provide Flowerheads are borne in small compact cypes with good sheep feed. In California small amounts of low densely pubescent branches. Involucral bracts are rabbitbrush were found in deer stomachs examined between October and January (Sampson and Jespersen 5.9 to 6.5 mm long, lanceolate to oblong, obtuse, and 1963). Much variation exists in palatability among the different subspecies. Some may be heavily utilized, whereas others are consumed little if at all (McArthur and others 1974). Variety stenophyllus of ssp. viscidiflorus on rocky foothills is often heavily used, and sometimes destructively so, with animals prefer- ring mature or partially mature plants to green immature ones. On a Utah winter range, this subspecies averaged to 11.31 percent by weight of the diet of sheep (Cook and Harris 1950; Cook and others 1954). On another Utah winter range, mule deer diets con- sisted of over 80 percent low rabbitbrush in December (Austin and Urness 1983). Substantial use of ssp. lanceolatus in widely scattered areas of Utah and Nevada has been observed (McArthur and others 1979a). Antelope, elk, and bighorn sheep, as well as deer and livestock, show varying preferences for low rabbitbrush, depending on season, locality, and subspecies B (Kufeld 1973; Kufeld and others 1973). The species, like rubber rabbitbrush, increases rapidly and vigor- D ously on otherwise disturbed sites. Some subspecies C such as stickyleaf low rabbitbrush (ssp. viscidiflorus) and mountain low rabbitbrush (ssp. lanceolatus) adapt well to higher elevations, while other subspecies such A as hairy low rabbitbrush (ssp. puberlus) do best in lower desert and foothill habitats. Low rabbitbrush is Figure 44—Three subspecies and a variety of valuable for revegetating depleted rangelands and low rabbitbrush: (A) C. viscidiflorus ssp. other disturbed sites such as strip mines and road- lanceolatus; (B) C. viscidiflorus ssp. publerus; sides (Plummer 1977). (C) C. viscidiflorus ssp. viscidiflorus var. stenophyllus; (D) C. viscidiflorus ssp. viscidiflorus.

532 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs glabrous to pubescent. Achenes are densely strigose. primarily in sagebrush and pinyon-juniper communi- On the basis of his systematic investigations in the ties at elevations between 5,000 and 8,500 ft (1,500 and genus Chrysothamnus, Anderson (1980) recom- 2,600 m). This subspecies often becomes dominant in mended placing the former ssp. elegans in synonymy cleared or overgrazed areas. At lower elevations with C. viscidiflorus ssp. lanceolatus. Chromatographic stickyleaf low rabbitbrush may be associated with work by McArthur and others (1978a) supports this such halophytes as shadscale, winterfat, and haloge- consolidation. Mountain low rabbitbrush is wide- ton. Other subspecies of low rabbitbrush are also spread and fairly common in dry foothill and moun- often associated with it. tainous habitats from 5,000 to 10,500 ft (1,500 to Anderson (1971) recommends placing the former 3,200 m) ranging from British Columbia, east to ssp. pumilis in synonymy with C. viscidiflorus ssp. Montana, and south to New Mexico, Utah, and Nevada. viscidiflorus because its specimens are “only environ- This subspecies may be found growing with such mentally modified variants of C. v. ssp. viscidiflorus.” shrubs as big sagebrush, snakeweed, various sub- Chromatographic work (McArthur and others 1978a) species of rubber rabbitbrush, Greene’s rabbitbrush supports Anderson’s reduction of pumilus to synonymy. (C. greenei) and Parry rabbitbrush (C. parryi) Because of its distribution from moist to arid sites, the (McArthur and others 1979a). species is well suited to a wide range of disturbed sites Hairy low rabbitbrush (Chrysothamnus viscidi- over the Western States (McArthur and others 1979a). florus ssp. puberlus) is a small shrub up to 20 inches Recently Anderson (1980) reduced ssp. stenophyllus (50.8 cm) high with yellowish to green, finely pubes- to varietal status under ssp. viscidiflorus. This variety cent branches (fig. 44). Its linear-filiform to linear is known as narrowleaf low rabbitbrush (fig. 44). It is leaves are sparsely to densely pubescent with scabrid- a low, glabrous shrub up to 12 inches (30.5 cm) high ciliate margins and are usually twisted or revolute. with white bark. Leaves are linear-filiform, often The leaves are up to 2 mm wide and up to 3 cm long. twisted, viscidulous, 1 mm or less wide, 1 to 3 cm long, Flowerheads are borne in small compact cymes with and glabrous except for the usually scabrid and revo- densely pubescent branches. Involucral bracts are lute margins. The branches of the small, compact about 5 to 6 mm long, lanceolate to oblong, acute to cymes are glabrous. Involucral bracts are 4 to 6 cm obtuse, and are usually marked with a thickened long, not keeled, and lance-oblong. greenish spot near their tips. Hairy low rabbitbrush Narrowleaf low rabbitbrush is rather common on occurs on dry plains, valleys, and foothills, especially many desert ranges, particularly in the northern on poorer soils and disturbed areas. Its range is essen- Great Basin and adjacent areas (Anderson 1980, tially the Great Basin area from southern Idaho and 1986c). On these ranges this variety is usually found Oregon south through Utah and Nevada to eastern in the sagebrush type on poorer soils and disturbed California and northern Arizona (Anderson 1986a). sites, but is also found growing with halophytes such This subspecies is most abundant in the big sagebrush as shadscale, fourwing saltbush, greasewood, and communities of western Utah, Nevada, and south- halogeton. central Idaho. However, it has been found growing in Anderson (1980, 1986a) recognized two other sub- one locality or another with most of the other subspe- species of low rabbitbrush. These are ssp. axillaris and cies of low rabbitbrush, shadscale, winterfat, haloge- planifolius. The former subspecies is morphologically ton, and occasionally with pinyon and juniper quite similar to var. stenophyllus, but occurs mostly (McArthur and others 1979a). in the southern Great Basin and on the Colorado Stickyleaf low rabbitbrush (Chrysothamnus viscidi- Plateau and appears by its somewhat attenuate florus ssp. viscidiflorus) is the largest subspecies of bracts to be introgressed by Greene’s rabbitbrush. It is low rabbitbrush. Mature shrubs are usually more quite abundant in certain locales. Subspecies than 20 inches (50.8 cm) tall, whereas the other sub- planifolius is a local form with small heads and flat species are normally under 20 inches (50.8 cm). Its leaves endemic to Coconino County, AZ (Anderson branches, leaves, and inflorescences are glabrous but 1986a). viscid (sticky). The broadly linear to narrowly lanceolate, bright green leaves are 1 to 5 mm wide, 2 to 5 cm long, and flat to twisted (fig. 44). Leaf margins are Other Rabbitbrushes ______sometimes scabrid. Crushed foliage usually emits a Alkali rabbitbrush (C. albidus) and Greene’s rabbit- pungent odor. Branches of the stems are glabrous. brush (C. greenei) are two other relatively important Involucral bracts are obtuse, oblong, not keeled, and species. Alakli rabbitbrush is a much branched, leafy 5 to 7 mm long. Stickyleaf low rabbitbrush is widely shrub. It has erect, brittle, glabrous, very resinous, distributed on dry plains and hills from Washington, whitebarked branches up to 3 ft (91.4 cm) high. The Idaho, Montana south to Colorado, Utah, Nevada, glabrous, filiform leaves are 1.5 to 4 cm long, 0.5 to northern Arizona, and eastern California. It occurs 2 mm wide, and the margins become revolute. Their

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 533 Chapter 21 Composite Shrubs surface is covered with small pits and abundant resin- on plains, valley, and foothills in Colorado, New Mexico, ous exudate. Heads with four to six white disc flowers Nevada, and Utah. Overgrazing allows it to greatly each are arranged in small compact cymes. Each head increase, sometimes forming a subclimax community is subtended by approximately 15 glabrous, resinous, (Hall and Clements 1923). This species provides cover involucral bracts. These are 7 to 9 mm long and and emergency browse in areas where it is abundant. terminate in attenuate to acuminate, usually curved There are several other rabbitbrush species tips. The pappus is abundant and longer than the (Anderson 1986a). These are all less common than corollas. Mature achenes are about 4 mm long and the species discussed or are outside of the geo- densely covered with long soft hairs. Blooming occurs graphical area of concern for this treatment. These from August to November. This species is a definite are C. gramineus and C. eremobius both related to the halophyte. It occurs most commonly along the western genus Petradoria and of limited distribution in up- side of the Great Salt Lake desert but is also found lands of southern Nevada (and for C. gramineus, across Nevada to eastern central California in alka- adjacent California). Chrysothamnus molestus is a line soils. Alkali rabbitbrush may invade strongly rare, distinctive species from Coconino County, AZ. alkaline areas as a pioneer plant. In less alkaline flats Both C. pulchellus and C. spatulatus are distinctive it may be associated with threadleaf rubber rabbit- species that occur southeast of the main distribution of brush, basin wildrye, and greasewood. This species the genus Chrysothamnus. Chrysothamnus humilus has value as ground cover on alkaline soils (McArthur is similar in some respects to low rabbitbrush, but is and others 1979a). distributed on the northwest fringe of the species distri- Greene’s rabbitbrush is a small highly branched butional range. Both C. paniculatus and C. teretifolius shrub only 4 to 14 inches (10.2 to 35.6 cm) high. Its are viscid hot desert southwest shrubs. The former is glabrous brittle twigs are green at first but soon more common in gravelly washes whereas the latter become white and shiny. Bark on the lower branches occurs mostly on rocky slopes. often peels off in sheets (Hall and Clements 1923). The nearly glabrous or slightly scabrous-ciliate leaves are narrow, linear, 1.2 cm or less wide, 1 to 3.5 cm long, and Other Composite Shrubs ______are more or less viscidulous. Flowerheads normally Haplopappus species (goldenweed), contain five disc flowers in rounded or flat topped Tetradymia species (horsebrush), cymes. The involucral bracts are 5 to 7 mm long, arranged in five poorly defined vertical ranks, and Gutierrezia species (matchbrush or terminate in narrowly acuminate, greenish tips. The snakeweed), Lepidospartum tubular to funnelform corollas may be whitish or latisquamatum (scalebroom) yellow and 4 to 4.5 mm long. Achenes are about 3 mm long and are covered with dense, long, shaggy hairs. Both goldenweed and snakeweed belong to the Greene’s rabbitbrush has been divided into two sub- same tribe (Astereae) of Compositae that rabbitbrush species, ssp. greenei and ssp. filifolius by some au- does (McArthur and others 1978a). Horsebrush and thors (Hall and Clements 1923; Harrington 1964; scalebroom belong to the tribe Senecionieae. Haplopap- Kearney and Peebles 1960). These authors separate pus is a large genus that is mostly herbaceous. Its ssp. filifolius from greenei by its normally larger stat- constituent species are widely distributed but usually ure and shorter narrower leaves. McArthur and others scattered over Western rangelands (Hall 1928; USDA (1978a) believe no subspecies should be recognized Forest Service 1937). The most notable shrubs are because the purported subspecies are (1) chromato- goldenweed rabbitbrush (Haplopappus bloomeri) graphically similar and (2) often occur in mixed popu- and its near relatives (Anderson 1983). Goldenweeds lations. The chromatographic data from McArthur are generally of little forage value, but they are of and others (1978a) further suggest that C. greenei value in erosion control (USDA Forest Service 1937). could be considered a subspecies of C. viscidiflorus Other shrubby goldenweeds include H. macronema, rather than a separate species. Chrysothamnus greenei H. suffruticosus, H. resinosus, H. greenei, and H. closely resembles C. viscidiflorus, particularly ssp. carthamoides (Hitchcock and others 1955). axillaris and ssp. viscidiflorus var. stenophyllus. All The genus Tetradymia consists of rather low, stiffly have a low bushy habit, whitebarked stems, and short branched shrubs (fig. 45). The stems are uniformly narrow leaves. Furthermore, ssp. axillaris has involu- canescent or have glabrous to woolly streaks running cral bracts with attenuate tips (Anderson 1964) that down the stem internodes from the primary leaves. closely resemble those of C. greenei. We concur with The tomentum may be permanent or deciduous on Hall and Clements (1923) who felt the similarity both the stem and leaves. Spines may be present between C. greenei and C. viscidiflorus indicated a (fig. 46). Tetradymia bears primary and secondary close genetic relationship. Greene’s rabbitbrush occurs leaves. Primary leaves develop alternately along

534 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs

elongated shoots and are usually long-lived. Second- ary leaves develop in fascicles in the axils of the primary leaves. They are generally short-lived and often dry up and fall away within a few weeks (Strother 1974). Horsebrush flowers are borne in heads located singly or in pairs in the upper primary leaf or spine axils or are clustered in short, dense racemes or corymbs at the tips of branches. Each head contains from four to nine yellow disc flowers. Ray flowers are lacking. Four to six equal involucral bracts with overlapping sides subtend each head. The pappus consists of numerous bristles or scales or may be lacking. The ovary of each flower develops into a glabrous to densely longhaired achene. This genus blooms from April to August depending Figure 45— Gray horsebrush growing at Wasatch on elevation, climatic conditions, and species. Numer- Station, Summit County, UT. ous small moths, bees, flies, and beetles visit the

A B C E D

F G

H J 5 cm I

Figure 46—Horsebrush species (Strother 1974): (A) Tetradymia canescens; (B) T. filifolia; (C) T. glabrata; (D) T. nuttallii; (E) T. stenolepis; (F) T. argyraea; (G) T. tetrameres; (H) T. comosa; (I) T. spinosa; (J) T. axillaris var. longispina.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 535 Chapter 21 Composite Shrubs flowers. Although numerous potential pollinators are available and the flowers are highly fertile, seedlings are not commonly seen in nature. This is probably due to the harsh environment in which horsebrush is usually found (Strother 1974). However, young plants and sprouts may be found growing in burned-over areas (McArthur and others 1979a). Horsebrush provides some critically needed ground cover in the dry, sparsely vegetated desert ranges where it grows. Al- though several species are poisonous to sheep and have caused losses of thousands of animals in Utah and Nevada, horsebrush is browsed particularly heavily during winter and early spring on desert ranges and overgrazed areas where little else may be available this time of year. In fact, Zimmerman (1980) Figure 47—Broom snakeweed, right, growing has built a successful winter range cattle operation in next to hairy low rabbitbrush, left, near Ephraim, central Nevada on horsebrush and other desert shrub Sanpete County, UT. species that are often considered useless. Severe losses have occurred when hungry animals have been trailed from winter to summer ranges through stands of horsebrush without allowing the animals a chance to graze other plants (Kingsbury 1964). Considerable generally pubescent. Blooming occurs from May to variation in toxicity within and between species of October. The numerous common names of this genus Tetradymia has been noted by Johnson (1974a,b), who are perhaps indicative of its wide distribution. The believed ingesting black sagebrush and horsebrush genus consists of about 25 species scattered through- under certain poorly understood conditions is respon- out western North and South America (Hitchcock and sible for toxicity. Four horsebrush species (T. canescens, others 1955; Lane 1985). Only two of these species, T. glabrata, T. nuttallii, and T. spinosa) are quite G. sarothrae and G. microcephala, are of importance common in the Intermountain area. The first two in the Intermountain area although there are two less species are spineless, and the latter two are spiny common endemics, G. petradoria and G. pomariensis (McArthur and others 1979a). Tetradymia axillaris, (Welsh and others 1987). Some additional North another spiny species, occurs on the edge of the Inter- American species include G. bracteata, G. californica, mountain area in scattered populations from south- and G. serotina (Lane 1985; McArthur and others western Utah to southern California. Two other spe- 1979a; Solbrig 1971; . Snakeweed commonly invades cies are found southwest of the Intermountain area: depleted ranges and is considered an indicator of T. argyraea (southeastern California) and T. comosa (southern California and Baja California) (McArthur and others 1979a; Strother 1974). The broom snakeweeds (Gutierrezia formerly Xanthocephalum) consist of mainly perennial herbs and low suffrutescent shrubs (subshrubs) with woody roots, crowns, and stem bases (fig. 47). A few species are annuals (Lane 1985). However, of these only, G. sphaerocephala occurs in the Intermountain area (in New Mexico). These perennials are ordinarily short-lived as well. Snakeweed leaves are entire, linear to narrowly oblanceolate, and usually sticky from resin exuded to the surfaces of both leaves and young stems. Numerous small heads form in loose or crowded terminal clusters. Resinous, imbricated involucral bracts within thin membranous margins and green tips subtend each head. A few yellow ray and disc flowers are both present (fig. 48). The ray flowers are usually pistillate and fertile, while the disc Figure 48—Broom snakeweed, right, with its flowers are usually perfect and fertile or sometimes showy ray flowers, growing next to Parry rabbit- staminate. Pappi of several small scales or awns are brush, left, in Salina Canyon, Sevier County, UT.

536 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 21 Composite Shrubs overgrazed rangelands. It does occur, usually in a low, This plant superficially resembles rubber rabbitbrush but fluctuating density, in ranges in good to excellent but is a member of the tribe Senecioneae and hence condition. Species in this genus are generally un- has diagnostic morphological as well as chemical dif- palatable and seldom grazed. When eaten in quantity, ferences from rabbitbrush (McArthur and others this plant is more or less poisonous to sheep and 1978a). Scalebroom has potential use as a revegeta- goats (Benson and Darrow 1945; Kearney and Peebles tion plant in areas where it is adapted (Graves and 1960). others 1975; Kay and others 1977e). These authors Scalebroom (Lepidospartum latisquamatum) is con- demonstrated that seed should be planted less than fined to sandy washes and gravelly plains in the 1 cm deep. Furthermore, seed viability is lost rapidly Mojave Desert extending to the southwestern portion (in less than 2 years) if not stored in sealed containers. of the Intermountain area (Kay and others 1977e).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 537 Chapter 21 Composite Shrubs

538 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Nancy L. Shaw Stephen B. Monsen Chapter Richard Stevens 22

Rosaceous Shrubs

Important shrubs of the Rose Family (Rosaceae) in the Inter- mountain region are distributed from blackbrush and salt desert shrub communities through high elevation forests and meadows. Growth habits of this group vary from trailing brambles to upright shrubs and small trees. Some species are evergreen while others are deciduous. Many of these species are highly valued for the cover, fruits, and forage they provide for wildlife and livestock. Rosaceous species that develop spreading root systems, root suckers, rhizomes, or stem layers provide soil stabilization. Several species are capable of regenerating following fire. Those that produce fragrant flowers or colorful fall foliage are prized for their ornamental value. Because of their browse value, antelope bitterbrush and several other rosaceous shrubs were among the first species to be used in wildlife habitat improvement efforts. Members of the bitterbrush-cliffrose complex as well as serviceberries, chokecherries, and the mountain mahoganies lend themselves to artificial seeding; their seed are fairly large and relatively easy to collect, clean, store, and plant. Additional species of this family are seeing increased use in response to the growing emphasis on use of native species for wildland revegetation and low maintenance landscaping, community restoration issues, mitigation for en- dangered species, and a general shift to employ revegetation when deemed necessary to conserve or restore ecosystem diversity and functionality.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 539 Chapter 22 Rosaceous Shrubs

Chapter Contents by Species

Amelanchier alnifolia (Saskatoon serviceberry) ...... 541 Amelanchier utahensis (Utah serviceberry) ...... 543 Cercocarpus betuloides (Birchleaf mountain mahogany) ...... 544 Cercocarpus intricatus (Littleleaf mountain mahogany) ...... 545 Cercocarpus ledifolius (Curlleaf mountain mahogany) ...... 545 Cercocarpus montanus (True mountain mahogany)...... 550 Coleogyne ramosissima (Blackbrush) ...... 552 Cotoneaster acutifolia (Peking cotoneaster) ...... 555 Cowania stansburiana (Stansbury cliffrose or cliffrose) ...... 556 Crataegus douglasii (Douglas or river hawthorn) ...... 560 Fallugia paradoxa (Apache plume) ...... 561 Holodiscus discolor (Creambush oceanspray) ...... 563 Holodiscus dumosus (Rockspirea) ...... 565 Peraphyllum ramosissimum (Indian apple) ...... 566 Physocarpus malvaceous (Mallowleaf ninebark) ...... 567 Potentilla fruticosa (Shrubby cinquefoil) ...... 569 Prunus americana (American plum) ...... 571 Prunus andersonii (Desert almond or Anderson peachbrush) ...... 572 Prunus besseyi (Bessey cherry) ...... 573 Prunus emarginata (Bitter cherry) ...... 574 Prunus fasciculata (Desert peachbrush) ...... 575 Prunus spinosa (Blackthorn) ...... 577 Prunus tomentosa (Nanking cherry) ...... 577 Prunus virginiana (Chokecherry) ...... 577 Purshia glandulosa (Desert bitterbrush) ...... 579 Purshia tridentata (Antelope bitterbrush) ...... 581 Rosa woodsii ultramontana (Woods rose) ...... 586 Rubus leucodermis (Black raspberry, blackcap) ...... 590 Rubus parviflorus (Thimbleberry) ...... 591 Sorbus scopulina (Greene’s mountain ash) ...... 592 Sorbus sitchensis (Sitka mountain ash) ...... 594 Spiraea betulifolia (Bridal wreath) ...... 594 Spiraea densiflora (Subalpine spiraea) ...... 596 Spiraea douglasii (Douglas spiraea)...... 596

Use of rosaceous species often presents challenges Seedlings of native or introduced species of Prunus to those involved in revegetation efforts. Size and and Rosa are traditionally grown by many nurseries quality of seed crops vary considerably from year to for conservation plantings. Antelope bitterbrush, Sas- year. Complex germination requirements, low initial katoon serviceberry, and curlleaf mountain mahogany growth rates, sensitivity to competition, and rodent or true mountain mahogany seedlings are commonly and bird preference for seed and seedlings of some grown for wildlife habitat improvement. Seedlings of species hinder their establishment and reduce sur- specific ecotypes and most other species must gener- vival. Appropriate planting practices and manage- ally be obtained through contract production. ment of young seedlings or plants are required to Characteristics of the site of origin of seed or plant- minimize these problems. Control of browsing by live- ing stock should be matched to those of the planting stock and wildlife during the first 2 to 3 years is often site as closely as possible; ecotypes of individual spe- essential to improve establishment and early growth. cies vary considerably in range of adaptability. Al- Bareroot or containerized planting stock is commonly though few rosaceous shrub cultivars have been se- used to shorten the period of seedling vulnerability. lected and released for commercial production of seed

540 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs and planting stock by the USDA Soil Conservation Although different geographic races have not been Service (Hassell 1988), a number of others are under identified, they are expected to occur within these study. Site requirements and characteristics of re- widely distributed species (Brinkman 1974a). In addi- leased cultivars have been extensively studied and tion, several hybrids have been reported (Cruise 1964). documented. Publications are available describing the Saskatoon serviceberry is distributed from Alaska attributes and site requirements of commonly col- and the Yukon east to Hudson’s Bay and south to lected antelope bitterbrush populations (Tiedemann California, New Mexico, and Nebraska (Harrington and Johnson 1983). Verification of seed origin of 1964; Welsh and others 1987). Plants occur from near wildland collections may be accomplished by monitor- sea level to over 9,000 ft (2,770 m) in the Rocky ing contract seed collections from designated native Mountains in areas with an annual precipitation of sites or plantings. Seed certifying agencies have devel- 12 to 25 inches (30 to 64 cm) or more (Jones 1946). oped a “source identified” classification for wildland Saskatoon serviceberry grows on sites ranging from seed certification that provides verification of site of dry, rocky, exposed slopes to riparian areas with deep origin as well as compliance with minimum seed moist soils. At the lower edge of its range, plants are quality standards (Young 1995). often intermixed with mountain big sagebrush/grass Seeding recommendations for major vegetative types and to a lesser extent with Wyoming big sagebrush/ and conditions are discussed in chapter 17. Rosaceous grass communities (Tisdale and Hironaka 1981). shrubs adapted to these vegetative types and condi- Saskatoon serviceberry also occurs in mountain brush, tions are included in the seeding recommendations. upper pinyon-juniper, and ponderosa pine communi- Seed characteristics are found in chapter 24. ties (Blauer and others 1975). It sometimes grows in The following sections include a brief species de- nearly pure stands, dominating other shrubs, but scription and discussions of ecological relationships, supporting a productive understory. distribution, culture requirements, use, improved varieties, and management for each of these Rosaceae Plant Culture species. A few small nurseries or conservation plantings of Saskatoon serviceberry are managed to furnish seed Amelanchier alnifolia ______for planting stock production. Most seed currently used in artificial revegetation is harvested from wild- Saskatoon Serviceberry land stands. Large quantities of seed can be harvested Saskatoon serviceberry is a deciduous shrub or at reasonable costs during good production years. small tree 1 to 20 ft (0.3 to 6 m) or more in height Some stands produce consistently good seed crops, but (fig. 1). The root system varies from a strong taproot harvestable yields usually occur at intervals ranging to a branching lateral root system. New shoots develop from 1 to 5 years, depending on the site. Plants from rhizomes extending up to 5 ft (1.5 m) from the usually produce numerous flowers each year, but parent plant (Hemmer 1975). Branches are reddish brown and pubescent when young, but later became glabrous. Leaf shape is extremely variable. They are approximately 1 inch (2.5 cm) long and oval to suborbicular with serrate or dentate margins to the middle or below. Perfect flowers with five showy white petals appear in large terminal clusters. Mature pomes are purple to black and 0.4 to 0.6 inch (1.0 to 1.5 cm) in diameter. Each pome contains 10 locules, each with one seed; one third to one half are commonly abortive (Blauer and others 1975; Brinkman 1974a; Harrington 1964). Seed are dark purplish-brown with a leathery seed coat (Belcher 1985). Flowers bloom from May through June and fruits ripen in July and August (Dittberner and Olson 1983).

Ecological Relationships and Distribution Figure 1—Saskatoon serviceberry occurs as Three species of serviceberry are distributed through single plants, thickets, and nearly pure stands on the western United States. Saskatoon serviceberry, sites ranging from riparian areas to dry, rocky dwarf Saskatoon serviceberry, and Utah serviceberry. slopes.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 541 Chapter 22 Rosaceous Shrubs spring frosts, rust infestations, and insect damage 1957). Komissarov (1938) found rooting occurred in often diminish seed production (Blauer and others 25 percent of all softwood cuttings taken in mid- 1975). Seed is disseminated by gravity and by birds summer following treatment with 50 mg/L indole-3- and other animals that pass the seed through their acetic acid (IAA) for 24 hours. Untreated cuttings did digestive tracts (Hemmer 1975). not root. Field survival of planting stock depends on Pomes are fleshy when mature and dry slowly on the use of healthy dormant or properly hardened stock shrub. They are normally collected by hand picking or and reduction of competition on the planting site. by beating the shrub to dislodge them. Seed can easily be separated from the fresh pomes. However, once the Uses and Management fruits dry, the pericarp remains closely attached to the seed. Dry pomes are macerated in a blender or Dybvig Saskatoon serviceberry is an important browse to separate the seed from the dried pulp. Dried material plant for game and livestock due to its fair to high is then extracted from the seed by screening. For seed palatability (Blauer and others 1975). It is capable of purchases recommended purity is 95 percent and persisting with a diverse and productive understory germination 85 percent. There are about 45,395 seed that is important for summer grazing. Saskatoon per lb (100,078 per kg) at 100 percent purity (see serviceberry is used by livestock, particularly sheep, chapter 24). Seeds have varying degrees of embryo in midsummer after full leaf development and after dormancy that can sometimes be overcome by wet more palatable species have been browsed (Blauer and prechilling (Brinkman 1974a). Seed prechilled at 34 others 1975). to 41 ∞F (1 to 5 ∞C) for 3 to 20 months germinated from 84 to 100 percent; the time requirement varied consid- Big game animals make considerable use of the erably among seedlots (Hargrave 1937; Heit 1968b; shrub (Kufeld 1973; Kufeld and others 1973). Tueller McKeever 1938). The seed coats of some serviceberry (1979) found serviceberry was utilized by deer during species also prevent germination (Young and Young the entire year, but it was most important during the 1986). Weber and others (1982) found acid scarifica- midsummer and fall months. New shoots are particu- tion for 30 minutes and imbibition in a mixture of larly palatable, but deer consuming large quantities of thiourea and benzyladenine released this form of dor- new serviceberry growth in spring can be poisoned by mancy. These authors found no differences in germi- it. Problems also occur if Saskatoon serviceberry com- nation among different sized seed. Some selections prises more than 35 percent of the fall or winter diet of from central Montana appear less dormant than many stressed deer (Quinton 1985). Birds utilize the bark, other populations. fruits, and leaves. The fruit is a key grizzly bear food Saskatoon serviceberry should be seeded separately (Mace and Bissell 1986; Zager 1980). Seed are used by from rapidly growing herbs. Plantings should be made rodents and small birds (Blauer and others 1975). in strips or scalps at least 25 to 30 inches (63 to 76 cm) Saskatoon serviceberry is particularly valuable as a wide. Seed may be drilled or broadcast seeded in a minor species in sagebrush communities on open hill- firm, prepared seedbed and should be covered with sides and ridges. Use should not be allowed to reduce 0.25 to 0.50 inch (0.64 to 1.25 cm) of soil. Broadcast plant productivity at these locations. seeding without soil coverage is not recommended. Saskatoon serviceberry is a useful shrub for land- Rodents quickly gather seed, even those placed in soil. scape plantings in recreation areas, roadways, and Consequently, seeding in late fall or early winter after other low maintenance areas. It persists in areas that rodent activity has diminished is advised (Brinkman receive heavy traffic and can withstand wind and 1974a; Plummer and others 1968) severe storms. It can also be used as a background Seedling growth is dramatically affected by avail- screen or for focal plantings. able soil water. Survival may be low if seedlings are Saskatoon serviceberry should not be planted off subjected to drought during the first few months of site. Sources from mountain brush communities are growth. Stressed plants are often stunted, and al- not well suited to lower elevations. The species has though they may persist through one or more growing not performed well on road or mine disturbances. seasons, they usually do not recover. In contrast, Developing seedlings do not survive if heavily grazed healthy seedlings that survive the first growing sea- or subjected to drought stress. However, established son are usually very tenacious. plants, those 5 or more years of age, can withstand Saskatoon serviceberry can be grown and field rather heavy use, particularly winter browsing. In planted as bareroot or container stock. Bareroot seed- fact, mature plants may be stimulated by some lings reach adequate size for outplanting in one sea- grazing. Plant vigor can be maintained with 60 per- son, developing a heavy taproot (Shaw 1984). Con- cent fall or winter use or 40 to 50 percent spring use tainer seedlings are propagated from seed, germinants, (Young and Payne 1948). or small transplants and require a 3 to 4 month Saskatoon serviceberry seedlings develop slower cropping period (Landis and Simonich 1984). The than antelope bitterbrush seedlings. Young seedlings shrub is not easily propagated from cuttings (Doran are seriously affected by grazing, and, unfortunately,

542 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs game animals tend to seek these plants. If not heavily short leafy branchlets. They are perfect and regular grazed, 4- to 5-year-old plants often begin to grow with five white petals. The stamens are numerous relatively rapidly. Such plants usually require 8 to 12 and inserted on the rim of the calyx tube. The ovary is years to reach mature size, but some plantings of inferior with three or four styles. Each locule con- adapted ecotypes established on dry sagebrush sites tains two ovules. Fruits are purple to black pomes have taken as much as 20 years to mature. Production 0.2 to 0.4 inch (6 to 10 mm) in diameter (Harrington of good seedcrops may not occur until shrubs reach 1964; Welsh and others 1987). full size. Moderate grazing of established plants may Utah serviceberry differs from Saskatoon service- keep bushes hedged, but this usually does not cause berry in having a somewhat smaller stature, smaller mortality. leaves that are pubescent at least on the lower surface, Mature plants tend to persist for long periods. three to four styles compared to four or five, and a Hemmer (1975) reported ages of 50 to 85 years for hard, dry fruit. Davis (1952) considered Utah service- serviceberry growing in climax brush fields. Saska- berry a subspecies of Saskatoon serviceberry, but toon serviceberry can remain suppressed in closed Jones (1946), Blauer and others (1975), and Welsh and conifers stands for long time periods. Reducing canopy others (1987) considered differences in fruiting and cover stimulates resprouting (Wright and others 1979). growth habits sufficient to warrant species status. Regeneration from seed is normally rare (Noste and Bushey 1987). Ecological Relationships and Distribution Saskatoon serviceberry is most vigorous in seral communities. Productivity may increase following Utah serviceberry occurs at scattered sites from burning or mechanical control in such situations. eastern Oregon to Montana and south to Texas, However, where density is low or succession far ad- California, Baja California and Sonora (Blauer and vanced, such efforts may be counterproductive others 1975; Welsh and others 1987). It is generally (Hemmer 1975; Howard 1997). Plants recover well found on drier sites than Saskatoon serviceberry. It following fires and survive even severe burns if the soil occurs on foothill sites at elevations from 2,000 to is moist at the time of burning (Fischer and Clayton 7,000 ft (600 to 2,600 m) where it grows on dry, rocky 1983). Tops may be killed, but plants can resprout outcrops and on shallow soils. It is associated with a from rhizomes (Frischknecht and Plummer 1955). wide number of plant communities including aspen, Those rhizome buds located immediately below the ponderosa pine, pinyon-juniper, mountain brush, burned tissue sprout most readily (Bradley 1984). and big sagebrush. Although it may occur in dense stands, plants are usually scattered and intermixed Varieties and Ecotypes with other woody species. Selection work has focused on comparing seed germination characteristics and site requirements of selected ecotypes. To date, studies have not resulted in identification of ecotypes unusually well suited to restoration and revegetation work, although some populations seem to produce consistently higher seed fill. For this reason, use of local material on sites previously occupied by Saskatoon serviceberry is recommended.

Amelanchier utahensis ______Utah Serviceberry Utah serviceberry is similar in growth habit to Saskatoon serviceberry. Plants vary from large upright shrubs to small trees 4 to 15 ft (1.2 to 4.6 m) tall (fig. 2). Twigs are unarmed. The alternate deciduous leaves are simple, about 0.8 to 2.0 inches (2 to 5 cm) long, and are borne on a short petiole (Harrington 1964). Leaves are suborbicular, oval, ovate or obovate Figure 2—Utah serviceberry is a highly vari- with coarsely serrate-dentate margins to the middle able species that usually occurs on drier sites or below. Flowers are clustered on racemes borne on than Saskatoon serviceberry.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 543 Chapter 22 Rosaceous Shrubs

Plant Culture protection from grazing until plants have fully established, but mature shrubs withstand considerable Fruits of Utah serviceberry ripen in late summer, browsing. Some early plantings have been productive but may remain on the bush for up to 2 years. Utah for 40 years, although little spread has been noted serviceberry plants generally produce fewer fruits (Monsen n.d.). Neither serviceberry species estab- than Saskatoon serviceberries; this may be due to its lishes or grows well on highly disturbed sites such as occurrence on drier sites. However, some stands in roadways or mine spoils. southern Utah’s Iron County often yield extremely Utah serviceberry provides good forage for cattle heavy crops. Seed are dispersed by birds and small and good to excellent browse for sheep and goats in mammals (Brinkman 1974a). early spring. Deer browse it year long, but use is The pomes are somewhat more difficult to dislodge particularly heavy in winter. Birds and small mam- than those of Saskatoon serviceberry but can be col- mals use the berries in winter and spring (Blauer and lected in fairly large quantities during years of good others 1975). Plants survive fires and regenerate by production. To obtain seed, the dry pomes are usually sprouting (Tisdale and Hironaka 1981; Wright and placed in a blender or a Dybvig with water and macer- others 1979). ated to separate the seed from the dry pulp. Seeds are smaller and often have lower fill than those of Varieties and Ecotypes Saskatoon serviceberry, but germination characteristics are quite similar. For seed pruchases, recommended There are no releases. Local ecotypes are recom- purity is 85 percent and germination 95 percent. mended for on-site planting. There are about 25,800 seeds per lb (56,900 per kg) at 100 percent purity (see chapter 24). Cercocarpus betuloides ______Utah serviceberry is best established by drill seeding. Seed should be planted at a depth of 0.5 inch Birchleaf Mountain Mahogany (1.3 cm) in a weed-free seedbed. Hand seeding disturbed sites or seeding with interseeders or similar Birchleaf mountain mahogany is an open, erect implements that can selectively place seed in the soil shrub or small tree varying in height from 6 to 23 ft are also effective. Seeding at rates of 2 to 4 lb per acre (2 to 7 m). Although evergreen, it is similar in other (2.2 to 4.4 kg per ha) is required to produce acceptable respects to true mountain mahogany (Welsh and stands. Seedlings establish slowly and are sensitive to others 1987) (fig. 3). It occupies dry slopes and washes herbaceous competition. Bareroot or container seed- below 6,000 ft (1,846 m) from central California to lings are reasonably easy to produce, but they are also southwestern Oregon (Munz and Keck 1959). highly sensitive to competition.

Uses and Management Utah serviceberry is adapted to low foothill sites, and may be used to improve diversity in sagebrush communities. Selections of Utah serviceberry have been seeded in relatively arid situations such as pinyon-juniper and big sagebrush communities where serviceberry is not native, in expectation that it may successfully establish and improve wildlife cover. Successful plantings in such areas have persisted for approximately 30 years to date, but natural recruitment has not occurred. Saskatoon serviceberry has failed to persist in such situations. Utah serviceberry is resistant to rust (Gymnosporangium spp.) diseases that often attack Saskatoon serviceberry (Krebill 1972; Plummer and others 1968) and is occasionally selected for planting where the rust is prevalent. Slow establishment and poor survival have somewhat limited the use of both serviceberry species, but Utah serviceberry establishes and grows faster than Figure 3—Birchleaf mountain mahogany Saskatoon serviceberry (Plummer and others 1968). stem with new leaves Seedlings fail to establish when seeded in poorly and flowers, just after prepared, weedy seedbeds. Young seedlings require petal fall.

544 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

Collections of birchleaf mountain mahogany from Cercocarpus ledifolius ______central California have been planted within the Intermountain region on game and livestock ranges Curlleaf Mountain Mahogany because of their evergreen growth habit, seedling vigor, and drought tolerance. Healthy birchleaf Curlleaf mountain mahogany is an erect shrub or mountain mahogany seedlings that survive the first small tree sometimes attaining heights of 22 to 26 ft growing season are usually very tenacious. However, (7 to 8 m) (fig. 5) (Harrington 1964; Munz and Keck young plants are often stunted when stressed and 1959). It has a rather deep, well developed root system although they may persist for 1 or more years, they (Noste and Bushey 1987) and one to several main usually do not recover. The seedlings and young plants trunks up to 7 inches (18 cm) in diameter with deeply grow slightly faster than most selections of true furrowed reddish-brown bark. The shiny leaves are mountain mahogany for about the first 5 or 6 years. thick and evergreen, 0.4 to 1.2 inches (1 to 3 cm) long After this time, growth rates and growth habits of the and 0.2 to 0.4 inch (0.5 to 1 cm) wide. Leaves are two species are quite similar. resinous and aromatic, entire, lanceolate, usually glabrous above and white tomentose below. Leaf margins are highly revolute, hence the name curlleaf. Cercocarpus intricatus ______Flowers are solitary or in twos or threes with white to greenish-yellow sepals and no petals. The fruit is an Littleleaf Mountain Mahogany achene 0.3 to 0.4 inch (0.8 to 1.0 cm) long with an Littleleaf mountain mahogany is an intricately attached plumose style elongating to 1.6 to 2.8 inches branched shrub usually less than 8 ft (2.4 m) tall (4 to 7 cm) at maturity. The flowering period extends (fig. 4), with strongly revolute linear leaves (Blauer from May to June; fruits ripen from May to August and others 1975). It occurs primarily in Utah, Nevada, (Blauer and others 1975; Harrington 1964; Munz and and southern Arizona, occupying harsh rocky sites, Keck 1959). particularly areas with high summer temperatures and infrequent rain storms. There are low-growing forms of littleleaf mountain mahogany in Oregon and Washington (Hitchcock and others 1961). Stutz (1974) suggests that littleleaf mountain mahogany is a seg- regant of curlleaf mountain mahogany that tends to be associated with more xeric sites. Seed germination and cultural requirements for littleleaf mountain mahogany and other mountain mahogany species are quite similar. Littleleaf mountain mahogany is productive and evergreen with a low shrubby growth habit. It hybridizes with both curlleaf and true mountain mahogany (Blauer and others 1975; Stutz 1974).

Figure 4—Littleleaf mountain mahogany is ever- Figure 5—Curlleaf mountain mahogany, note high- green, low-growing, and generally very palatable. lining by deer.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 545 Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution 500 per acre (200 per ha) in eastern Oregon. Somewhat higher numbers were reported from Montana, the Curlleaf mountain mahogany is distributed east of average figure being 1,375 stems per acre (550 per ha) the Cascades and Sierra Nevadas from Washington to (Duncan 1975). In Idaho, Scheldt (1969) reported a Baja California and east to Montana, Colorado, and mean of 207 shrubs per acre (84 per ha). Arizona. Curlleaf mountain mahogany occurs at ele- Three principal species of mountain mahogany vations from 2,000 to 9,000 ft (615 to 2,770 m) (Cercocarpus ledifolius, C. intricatus, and C. montanus) (Harrington 1964; USDA Forest Service 1937). occur in the Intermountain region (Harrington 1964; Curlleaf mountain mahogany is usually found on Welsh 1982). Holmgren (1987) divided C. ledifolius warm, dry, rocky ridges on southern or western slopes into two taxa: C. ledifolius var. ledifolius and var. but may occur on all exposures (Brayton and Mooney intermontanus (Intermountain curlleaf mountain 1966; Martin 1950; Miller 1964; Tidestrom 1925). At mahogany). Intermountain curlleaf mountain ma- lower elevations in the Great Basin, it often grows hogany is distinguished from variety ledifolius by its on limestone, decomposed granite, or other coarse- broader leaves that are less densely pubescent ven- textured soils on steep north slopes and among cliffs trally, its usually longer petioles and hypanthium and ledges (Dayton 1931). Plants may be nodulated by tubes, and more treelike habit. Stutz (1990) concurred the nitrogen-fixing endophyte Frankia (Lepper and with Holmgren’s revision, concluding that the taxa Fleschmen 1977). growing in the Intermountain West apparently ar- Curlleaf mountain mahogany characteristically rived from different origins. The evolutionary pro- grows in isolated patches that are nearly pure stands. cesses that produced the existing species, particularly These sometimes cover considerable acreages and littleleaf mountain mahogany are still proceeding. often form a band between conifer forests and sage- All three hybridize in areas where they overlap brush or other shrub communities. Curlleaf mountain (Blauer and others 1975; Plummer and others 1957; mahogany is not reported as a dominant or associated Pyrah 1964; Stutz 1974). The resulting hybrid popula- species of principal forest habitat types in Montana tions are frequently important components of local (Pfister and others 1977), northern Idaho, or vegetation. Washington (Daubenmire and Daubenmire 1968). Intermountain curlleaf mountain mahogany occurs However, the plant is a codominant with Douglas-fir in southeastern Washington, eastern Oregon, north- in some drier forested areas in Idaho (Steele and ern, eastern, and southern California, central and others 1981). Dealy and others (1981) described five southern Idaho, western Wyoming, Utah, and adjacent communities dominated by curlleaf mountain ma- parts of Colorado and Arizona (Holmgren 1987). It hogany in southeastern Oregon. These include curl- often exists in pure stands surrounded by open sage- leaf mountain mahogany/mountain big sagebrush/ brush or mixed with Gambel oak, bigtooth maple, bunchgrass, curlleaf mountain mahogany/mountain pinyon-juniper, ponderosa pine, Douglas-fir, or white snowberry/grass, curlleaf mountain mahogany/Idaho pine at elevations of 4,300 to 9,800 ft (1,300 to 3,000 m). fescue, curlleaf mountain mahogany/bearded Variety ledifolius occurs in eastern Oregon, central bluebunch wheatgrass, and curlleaf mountain ma- and southern Idaho, southwestern Montana, north- hogany/pinegrass. Johnson and Simon (1987) described a curlleaf mountain mahogany plant community type central Wyoming, and northern Utah (Holmgren 1987). for the Wallowa-Snake Province of northeastern It grows on talus slides, rock outcrops and rocky slopes Oregon. Curlleaf mountain mahogany is the dominant at elevations from 1,300 to 6,900 ft (400 to 2,100 m). understory in some limber pine, white fir, and juniper series (Dealy and others 1981; Hironaka and others Plant Culture 1983; Steele and others 1981; Youngblood and Mauk 1985). It also grows in association with other woody Stands of curlleaf mountain mahogany produce good species such as pinyon, oak, serviceberry, bitterbrush, seed crops about in 2 of every 5 years. Few years occur or gooseberry (USDA Forest Service 1937). Curlleaf when little or no seed develops. However, to be profit- mountain mahogany occasionally occurs with pon- able for hand harvesting, a heavy seed crop with good derosa pine where both species overlap, but these viability must be produced; seed must ripen fairly sites generally do not form large or distinct communi- uniformly; and most seed must remain on the bush ties. Extremely tall curlleaf mountain mahogany plants for a period of time to permit a collection period. occur with Douglas-fir and Rocky Mountain maple in Elevational range and aspect affect the period of canyon bottoms and narrow draws or drainages in flowering and seed development. Seed are sometimes central and southern Utah. These small communities dispersed rapidly by stormy weather conditions. Con- provide important habitat for big game and livestock. sequently, ripening seed should be checked periodi- In most situations where curlleaf mountain ma- cally if possible, and harvested immediately when hogany dominates, its crown cover ranges from 35 to mature. Ripe seed are normally wind dispersed and 60 percent. Dealy and others (1981) counted 300 to 800 can be carried some distance from the parent plant stems per acre (120 to 325 per ha) with an average of (Deitschman and others 1974a). Once the achenes fall

546 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs to the ground, rodents begin gathering them; a high Concentrating seedings in areas with low rodent percent of all viable seed may be removed within 1 or populations can greatly enhance seeding success. 2 days. Field mice remove embryos from the achenes. Curlleaf mountain mahogany seedlings are not Consequently, fallen achenes should be carefully ex- competitive with exotic annuals, particularly amined before they are collected. Twisting of the cheatgrass. Attempts to plant curlleaf mountain ma- plume in response to changes in water content plants hogany at lower elevations of mountain brush and big the seed by drilling it into the ground (Dealy 1978). sagebrush communities infested with these weeds Seed are harvested by beating the shrub to dislodge have generally not been successful. Proper seedbed the achenes and catching them in lightweight tarps or preparation and control of competing herbs are essen- netting placed down wind. Plumes can be removed tial for shrub establishment. Phillips (1970) reported with a Dybvig or barley debearder. The material is high seed germination occurred on mineral soils cleared then screened with a fanning mill to separate the seed of litter and plant competition. from the remaining debris. If harvested lots contain Seedlings emerge early in spring following ground numerous small twigs, they must be separated over a thaw when soil water is likely to be favorable. How- gravity table to improve purity. Seeds are not removed ever, seedlings are sensitive to late frosts. Dealy (1975) from the achenes. The ripened seed are hard and found new seedlings exhibit rapid root growth in brittle and can be fractured or broken by improper relation to top growth, providing some resistance to cleaning procedures, thus hammermilling should be drought and codeveloping competition. However, avoided. Purity of at least 90 percent and germination Scheldt and Tisdale (1970) did find heavy seedling of at least 80 percent are recommended for seed mortality occurred during the first summer of estab- purchases. There are about 51,865 seeds per lb lishment, often resulting from drought and predation (114,342 per kg) at 100 percent purity (see chapter by rodents and rabbits. 24). Curlleaf mountain mahogany seeds remain viable Curlleaf mountain mahogany can be propagated for extended periods in storage. Stevens and others and field planted as bareroot or container stock (1981a) found little loss of viability after 7 years of (Butterfield and Tueller 1980; Landis and Simonich warehouse storage. Nearly 85 percent of seeds stored 1984; Shaw 1984). Seedlings develop rapidly in bareroot for 15 years germinated. nurseries. They must be lifted when dormant. Ever- Freshly harvested seed is dormant and requires wet green plants are particularly sensitive to lifting and prechilling (Dealy 1975; Deitschman and others 1974a). handling once growth has begun. Damage to the tap- Dealy (1975) found seed prechilled at 39 ∞F (4 ∞C) for root should be avoided during lifting. Container stock 170 or 270 days germinated to 98 and 100 percent, is easier to handle and often provides better estab- respectively. He concluded that the membrane sur- lishment (Ferguson 1983; Landis and Simonich 1984). rounding the embryo may block gas diffusion and be a Propagation of curlleaf mountain mahogany from principal factor in controlling dormancy. Young and stem cuttings is difficult (Everett and others 1978). others (1978) found that soaking seed at 41 ∞F (5 ∞C) for 21 days in an aerated solution of potassium nitrate Uses and Management and gibberellin improved germination over a range of temperatures. Drying the seed narrowed the tempera- Because of its evergreen habit and occurrence be- ture range permitting germination, but increased tween forest and shrub communities, curlleaf mountain germinability was retained at these temperatures. mahogany provides important cover and forage for a Curlleaf mountain mahogany fruit size varies wide variety of wildlife species. Curlleaf mountain among ecotypes and collection sites. To date, seed size mahogany produces highly palatable winter browse has not been closely correlated with site conditions. and thermal and hiding cover for elk, deer and other Differences may be due, in part, to genetic traits; big game (Dealy 1971, 1975; Kufeld and others 1973; sources producing large fruits tend to perpetuate this Mueggler and Stewart 1980; Smith 1952). Tueller trait when planted on different sites. (1979) reported curlleaf mountain mahogany was one The oblong achenes of curlleaf mountain mahogany of six shrub species that comprised the largest single are large enough that they are easily handled and fraction of deer diets on winter ranges in Nevada. It is planted alone or in mixtures with other shrub seed also ranked as a principal winter browse for big game using conventional drills. They may also be hand in Utah (Richens 1967), Oregon, and Washington seeded in selected spots. Curlleaf mountain mahogany (Mitchell 1951). Curlleaf mountain mahogany pro- establishes best from fall seeding because of its wet vides food, hiding cover, and nesting sites for many prechilling requirement and ability to germinate at birds (Dealy and others 1981). low temperatures (Dealy 1975; Plummer and others Blauer and others (1975), Duncan (1975) and 1968). In addition, late fall or early winter seeding, Mueggler and Stewart (1980) reported livestock make when rodents are less active, reduces seed predation. light use of mature curlleaf mountain mahogany shrubs. However, seedlings and young stands can be

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 547 Chapter 22 Rosaceous Shrubs heavily browsed. Many healthy stands support a di- Plantings and stand rejuvenation treatments should verse herbaceous understory that provides good sum- be extensive to reduce the impact of browsing during mer forage for sheep and cattle (Dealy and others establishment or recovery (Gruell and others 1985). 1981), but they are often inaccessible to livestock in Plantings established in south central Idaho and pro- winter. Curlleaf mountain mahogany has good nutri- tected from livestock and deer use grew relatively tive value (5.9 percent digestible protein) and digest- slowly for about 6 years, but then grew dramatically ibility (64.8 percent) in winter (Bissell and Strong during the next 4 to 6 years. This growth pattern has 1955; Smith 1952; Welch 1981). It is one of only a few been noted in other protected study locations (Monsen shrubs to exceed the protein requirements of winter- n.d.). Because the species is highly palatable (Mueggler ing mule deer; its leaves are evergreen and retain high and Stewart 1980; Smith 1952), the naturally slow- protein levels throughout the winter (Welch and growing plants are continually browsed and, if not McArthur 1979b). Winter phosphorus level (0.18 per- protected, remain in a stunted condition, assuming a cent) exceeds the lower range of phosphorus required rounded, hedged form (Blauer and others 1975; Garri- for wintering mule deer (Trout and Thiessen 1973; son 1953). Plantings established in central Utah in Tueller 1979). 1950, for example, have survived continuous heavy Curlleaf mountain mahogany is a useful species browsing, and the shrubs have remained quite small. for revegetating roadway and mine disturbances Once released from grazing, hedged plants can recover (Hungerford 1984; Stark 1966) and it is an excellent quickly. Ferguson (1983) recommended browsing be ornamental. It requires little maintenance, the leaves limited to 50 to 60 percent of current annual growth to are attractive and evergreen, and it withstands con- maintain productivity and a vigorous, shrubby growth siderable traffic and browsing. Compared to other habit. wildland shrubs, it responds well to supplemental Many extensive stands of curlleaf mountain ma- watering. Curlleaf mountain mahogany wood is ex- hogany have been highlined. Most plants are mature tremely dense as a result of its low growth rate. It has to decadent. New growth is out of reach of browsing been used for charcoal, fenceposts, and wood carving animals. Growth from lower branches diminishes as (Blauer and others 1975). Stands are occasionally a result of hedging and shading. Little or no reproduc- logged for firewood. tion occurs in these closed stands (Phillips 1970). Curlleaf mountain mahogany exhibits three charac- Various treatments have been investigated to regen- teristics that should be recognized and understood if erate them. Stands with live branches within reach of the species is to be used in revegetation projects. First, browsing animals have been treated by top pruning in seedlings can perform well on exposed sites if pro- Utah and Montana. Pruning in spring or early fall tected from competition. Second, young plants can prompted regrowth and increased forage availability persist and furnish considerable high quality forage (Austin and Urness 1980; Garrison 1953; Ormiston even when moderately to heavily grazed. Under such 1978; Thompson 1970), but such treatments are ex- conditions plants will not reach mature stature and pensive. Chaining or bulldozing kills most older or could be weakened or killed if subjected to other forms highlined, single stemmed plants (Christensen and of stress. Third, although seedlings are not compatible others 1966; Dealy 1971), but can be used to open with herbaceous competition, established plants do areas within closed stands for natural regeneration or very well in conjunction with some highly productive planting. herbs. Curlleaf mountain mahogany could be more Plants growing on shallow soils and exposed sites, widely planted with some herbaceous species to im- usually midwinter ranges for game, are difficult to prove total and seasonal forage production if the shrubs manage. Curlleaf mountain mahogany growing in were planted in separate strips or spots from the monotypic stands or in mixed stands with other shrubs herbaceous species. such as antelope bitterbrush, skunkbush sumac, and Many stands of curlleaf mountain mahogany exist green ephedra often receives heavy browsing year with little herbaceous understory. Shrub/grass after year. Most of these species are capable of with- plantings established with intermediate wheatgrass standing considerable use, but may be weakened and and other introduced herbs can be very productive. killed over time. Natural recovery is slow and often However, the ability of the shrub to regenerate natu- prevented by continued use. Artificial revegetation is rally under such conditions is reduced, particularly on also very difficult on these sites. deep, productive soils. On shallow, rocky soils, where Curlleaf mountain mahogany plants are not highly weedy grasses and herbs have not gained dominance, fire tolerant. The plants and onsite seed banks can be native forbs can usually be seeded along with the killed by high intensity burns (Noste and Bushey shrubs. 1987; Wright and others 1979). The thick lower bark Curlleaf mountain mahogany requires a longer pe- protects plants from low-intensity fires and although riod to attain mature stature than most other shrubs. the species is a weak sprouter, some surviving plants

548 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs can resprout from stem buds. In addition, new seed- three species are closely related, each with chromo- lings may emerge from unburned seed (Bushey and some number 2n = 18 (McArthur and Sandersen Noste 1987; Gruell and others 1985). Some stands are 1985), artificial hybridization is possible and could be now restricted to rocky slopes due to high wildfire employed to incorporate specific traits into selected frequency. In other areas, intense fire suppression and populations. understory reduction with grazing have permitted Natural hybrids between Intermountain mountain curlleaf mountain mahogany to invade sites with mahogany and true mountain mahogany and later deeper soils (Dealy 1975; Gruell and others 1985). generation backcrosses are quite common where both Low intensity fires can be used to stimulate commu- parents are found growing together (Blauer and oth- nities with low vigor or maintain curlleaf mountain ers 1975; Plummer and others 1957). Hybrids are mahogany where it occurs as a seral species with somewhat fire tolerant, a characteristic of true conifers (Gruell and others 1985). Burning can also mountain mahogany. They often have a wider area of improve age class structure of curlleaf mountain ma- adaptation than curlleaf mountain mahogany. Hy- hogany stands. Natural recovery can occur following brids are easy to recognize; they are tall and treelike prescribed fires, but it is often significantly enhanced with evergreen leaves and always occur in stands of by mechanical site preparation that reduces herba- true mountain mahogany where true mountain ma- ceous competition and creates a weed free seedbed hogany is the female parent (Stutz 1990). These hy- (Phillips 1970). Stands may also be rejuvenated by brids rarely reproduce. Few hybrids of C. ledifolius thinning them to 40 to 60 clumps per acre (99 to var. ledifolius and true mountain mahogany have 148 per ha) and mechanically clearing seedbed areas been reported. This is perhaps due to the fact that within the stand. Protection of stands less than 50 these taxa occur together only infrequently and hy- years of age from fire is recommended if curlleaf mountain mahogany is the climax species. Reduction brids are less easily recognized. They are highly fer- of browse use is essential for all stand improvement tile. Hybrids between littleleaf mountain mahogany projects. and true mountain mahogany are rare (Pyrah 1964), The mountain mahoganies are relatively free from perhaps due to reproductive barriers (Stutz 1990). infestation by insects and disease (Ferguson 1983). They are usually sterile. Hybrids between littleleaf However, serious outbreaks have destroyed sizeable mountain mahogany and Intermountain mountain stands in southwestern Idaho. Furniss and Barr (1967) mahogany are common in some areas where the spe- reported the looper destroyed 46 percent of a 5,900 cies overlap (Blauer and others 1975; Plummer and acre (14,580 ha) stand of curlleaf mountain mahogany others 1957; Stutz 1990). over 3 years. Tent caterpillars have been responsible Cercocarpus ledifolius var. ledifolius is a small shrub for severe defoliation in the Great Basin (Furniss and or tree occurring in the northern part of the Intermoun- Barr 1967). tain region, and is most common in Wyoming and Montana (Cary 1917; Miller 1964). It also occurs in Varieties and Ecotypes Oregon, Idaho, California, Nevada, Utah, Colorado, and Arizona. It is taller in stature than littleleaf moun- Numerous accessions of curlleaf mountain mahogany tain mahogany with larger leaves and flowers. Hybrid- have been collected and evaluated for forage value, ization with curlleaf mountain mahogany is common. seedling establishment, and vegetative growth char- Some C. ledifolius var. ledifolius populations ex- acteristics. Attempts have been made to select and press unique features of value in range and wildlife develop ecotypes with improved seedling vigor and habitat improvement. Ecotypes occurring in eastern high growth rates, but to date no major improvements Wyoming and western Montana include low decum- have been achieved. In general, selections appear bent and semi-erect forms (Miller 1964). They are universally adapted to all areas where the species evergreen, propagate by stem layering, and produce occurs. However, it is possible that differences exist, profusely branching root systems (Miller 1964). Some but have not yet been fully recognized. plants are also capable of resprouting, a feature un- Differentiation of the ancestral mountain mahogany common in curlleaf mountain mahogany. The decum- may have resulted in the appearance of some impor- bent ecotypes exist primarily on shallow soils devel- tant traits. Stutz (1974) proposed that the highly oped from a fractured limestone substrate. plastic true mountain mahogany, a plant better adapted Members of these ecotypes vary in stature and to mesic conditions, may have evolved from curlleaf growth form. Most develop multiple stems and are mountain mahogany. In contrast, the smaller, nar- semi-erect with leaves of varying sizes. The decum- rower leaves of littleleaf mountain mahogany may bent growth form and the hybrids provide an ex- have evolved as a response of curlleaf mountain tremely valuable base for future plantings. If adapted mahogany to more xeric circumstances. Because the to typical curlleaf mountain mahogany sites, the

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 549 Chapter 22 Rosaceous Shrubs hybrids could be used to decrease the size of the tree- mahogany hybrids are most abundant because oftheir like curlleaf mountain mahogany growth forms. In overlapping flowering periods. True mountain ma- addition, the stem layering and root sprouting fea- hogany x curlleaf mountain mahogany hybrids are tures could enhance the fire tolerance and soil stabili- less common, and true mountain mahogany x littleleaf zation values of this important shrub. mountain mahogany hybrids are rare (Pyrah 1964). Ecological Relationships and Distribution Cercocarpus montanus ______The range of true mountain mahogany extends from True Mountain Mahogany Montana to South Dakota and south to New Mexico, Arizona, and Colorado (Harrington 1964; Martin 1950) True mountain mahogany also known as blackbrush, deerbrush, alder, sweetbriar, and tallow brush, is an in areas receiving 10 to 23 inches (25 to 58 cm) of erect shrub or small tree 2 to 13 ft (0.6 to 4 m) in height. annual precipitation. It occupies bluffs, mountain Stems of main branches have smooth, grayish-brown slopes, and foothills from 3,500 to 10,000 ft (1,070 to bark (fig. 6). Young twigs are often villous. The short 3,050 m) in elevation. It commonly grows on rocky or petiolate leaves are oval to broadly ovate, serrate well-drained soils on all exposures. It is also common above the middle, 0.4 to 2.0 inches (1 to 5 cm) long, and on moist fertile sites, canyon bottoms, and north and 0.4 to 1 inch (1 to 2.5 cm) wide. Leaves are deciduous east slopes. It usually grows in association with other to semi-evergreen, dark green and glabrous above shrubs and trees including ponderosa pine, Gambel with prominent veins and a fine whitish tomentum oak, Utah juniper, pinyon, and other woody species beneath. Flowers are perfect and regular, borne in of mountain brush communities (Blauer and others small clusters in the axils of short spurs. There are 1975). It frequently grows with mountain big sage- five sepals extending from a trumpetlike hy- brush. Pure stands do exist, but are limited in extent. panthium, no petals, 25 to 40 stamens in two to three In Utah the species is widespread, occupying diverse whorls, and a single pistil. The fruit is an elongate situations, but most commonly occurring on shallow achene with a plumose style, 2.4 to 3.9 inches (6 to soils and slickrock sites (Greenwood and Brotherson 10 cm) long at maturity. Flowering occurs from mid- 1978). In Colorado, Medin (1960) found soil depth to be May to late June. Fruits mature from July into Sep- the most important factor influencing true mountain tember depending on elevation. They are wind dis- mahogany production on both shale and sandstone- persed (Harrington 1964; Hitchcock and others 1961; derived soils. Clay content of the A horizon was very Welsh and others 1987). important in sandstone-derived soils; greater clay Chromosome number of true mountain mahogany is content was beneficial. Factors influencing water avail- n = 9 (McArthur and Sanderson 1985). Natural hy- ability to the plant were key in influencing production. bridization between species of mountain mahogany in Brotherson and others (1984) also found factors Utah has been observed in areas of overlapping ranges. Curlleaf mountain mahogany x littleleaf mountain affecting water availability had a significant impact on the development of true mountain mahogany communities. In Utah, Anderson (1974) reported a combination of site factors including soil texture, percent litter, soil pH, and exposed rock influenced the distribution of the shrub. However, no specific combinations of factors could be used to predict its occurrence. Stands on north-facing slopes of central Utah study sites were transitional in succession to other mountain brush types (Christensen 1964). Succession progressed much more slowly on more xeric southerly sites, and perhaps would never progress beyond the true mountain mahogany stage (Brotherson and others 1984).

Plant Culture Reproductive phenology of true mountain mahogany is related to aspect and elevation (Blauer and others 1975). Stands of true mountain mahogany growing on fertile sites usually produce an abundant seed crop. Figure 6—True mountain mahogany post-flowering, but prior to development of the elongate plu- Seeds are more easily harvested than those of curlleaf mose styles. mountain mahogany; plants are smaller in stature.

550 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

Seed is usually hand collected from wildland stands; all mahogany species is only fair to good. However, no commercial nurseries are being maintained. true mountain mahogany usually transplants better Plumes are removed with a Dybvig or barley debearder, than curlleaf mountain mahogany. followed by screening. Piatt (1973) reported that the larger seed of true mountain mahogany germinate Uses and Management better than small seed, but relative size may not be a reliable index of differences in source germinability. Medin and Anderson (1979) and Tueller (1979) re- Seed may be stored for up to 7 years in an open ported true mountain mahogany is one of the princi- warehouse without a major decrease in viability pal browse species used by wintering deer within its (Stevens and others 1981a). Purity of 90 percent and range. The widespread occurrence, nutritive value, germination of 80 percent are recommended for seed and high palatability of this species ensure its rank purchases. There are approximately 59,030 seeds per lb as a valuable shrub for big game habitat (Plummer (130,138 per kg) at 100 percent purity (see chapter 24). and others 1968; Young and Bailey 1975). The plant Ecotypes of this species appear much more sensi- also receives considerable spring and summer use by tive to site conditions than do collections of curlleaf big game and livestock and provides cover for numer- mountain mahogany. Utilization of ecotypes acquired ous wildlife species. Compared to curlleaf mountain near the proposed planting site or from similar sites is mahogany it is a superior browse for livestock, par- recommended. ticularly sheep (Ferguson 1983). True mountain ma- Seed must be fall sown at a depth of approximately hogany is fairly tolerant of browsing. A Colorado 0.5 inch (1.25 cm). Surface plantings are not suc- clipping study indicated plants can withstand 60 to cessful and should be avoided. Growth of emerging 80 percent browsing (Shepherd 1971). However, seedlings is seriously depressed if they are subjected Brotherson and others (1984) found that about one- to competition from herbs. Consequently, the shrub half of the true mountain mahogany in central Utah should be seeded in strips or plots separate from stands was highlined and out of reach of browsing grasses. Seeding this species with a Hansen seed animals. Some stands have been severely reduced by dribbler attached to a caterpillar tractor has been very heavy browsing (Ream 1964). successful on pinyon-juniper chainings. This method Nutrient value of true mountain mahogany in win- also serves to prevent development of such a dense ter is not as high as that of curlleaf mountain ma- shrub overstory that herbaceous species are elimi- hogany. Digestibility of true mountain mahogany nated. Interseeding the shrub into established stands has been reported as 48.4 percent. Digestible protein of perennial grasses can be successful if strips or is 3.4 percent (Smith 1957; Urness and others 1977; patches of understory vegetation are first removed. Welch 1981); adequate winter digestible protein con- True mountain mahogany is frequently scattered tent is 5.2 percent for wintering deer. among the upper big sagebrush communities. In those True mountain mahogany also has value as an situations it has sometimes been eliminated or seri- ornamental. Plants are used for landscaping along ously reduced by browsing. Natural recovery is slow, the West coast. In inland areas they are very hardy, and artificial seedings are difficult to establish if the requiring low maintenance. Its semi-evergreen habit, sites are infested with weedy annuals. If populations shiny plumose fruits, and growth habit are unique of exotic annuals are low, natural and artificial seedings and attractive. It grows rapidly with limited irrigation. are quite successful. Burning is a good technique for rejuvenating stands Germination requirements are similar to those of and increasing plant vigor. True mountain mahogany curlleaf mountain mahogany; seeds require after- can resprout vigorously from surviving root crowns ripening and wet prechilling (Deitschman and others following burning. Recovery may be rapid even fol- 1974a). Seedlings emerge shortly after the ground lowing rather intense burns (Blauer and others 1975; thaws. Many of the small seedlings are lost to drought Wright and others 1979). Survival after burning may and early spring frost (Plummer and others 1968). be high at all seasons, but response is greater follow- Mortality also results from severe browsing by ro- ing dormant season burns in Colorado (Young 1983). dents, rabbits, and big game. Consequently, seedings Current annual growth measured in late winter should be protected from browsing for 3 years or more. was 475 to 650 percent greater on burns than on Seedlings are easily grown as either container or controls. Production was greater on low intensity than bareroot stock. Planting stock must be dormant or on high intensity burns. Seed dispersed long distances properly hardened to ensure success. Bareroot seed- by winds from unburned sites also contributes to lings should not be root pruned during lifting or recovery of burned or otherwise disturbed sites. outplanting. Transplants should be planted in strips True mountain mahogany has been established or scalps cleared of competing vegetation. Field sur- from direct seedings on mine disturbances in Utah and vival of container or nursery grown transplants of Idaho (Ferguson and Frischknecht 1985). Seedlings

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 551 Chapter 22 Rosaceous Shrubs survive on infertile soils; on spoil wastes they appear stems proliferate by stem splitting. The short branches to benefit by the addition of even small amounts of die back from the apex after a few years of growth topsoil. Plants may be nodulated by the nitrogen- leaving a spiny tip. Lateral branches then develop fixing endophyte Frankia (Hoeppel and Wollum producing a rounded shrub (Bowns and West 1976). 1971). Transplants grow slowly and are sensitive to The small persistent leaves are clustered along the competition. branches. They are simple, leathery, about 0.2 to 0.6 inch (5 to 15 mm) long, and 0.04 to 0.12 inch (1 to Varieties and Ecotypes; Other Species 1.5 mm) wide. The flowers are terminal on new branchlets. They have four yellow to brownish sepals Attempts have been made to enhance seed germi- and are usually apetalous. Numerous stamens are nation, seedling vigor, and growth rate of true moun- inserted at the base of the disk that encircles the tain mahogany by selection of ecotypes. A selection ovary. The fruit is a glabrous, brown achene 0.2 to referred to as “Montane” was released by the U.S. 0.3 inch (5 and 8 mm) in length (Harrington 1964; Department of Agriculture, Natural Resource Conser- Munz and Keck 1959; Welsh and others 1987). Plants vation Service, Los Lunas Plant Materials Center normally flower as early as March at lower elevations (Thornburg 1982). This cultivar, collected on the Santa (McMinn 1951) and as late as June at higher eleva- Fe National Forest near Coyote, NM, has nondormant tions (Munz and Keck 1959). Seeds mature in mid- seed and excellent seedling vigor. Its adaptation to summer. Blackbrush has n = 8 chromosomes, which is sites outside its area of collection has not been fully unusual for the subfamily Rosoideae (McArthur and documented. Sanderson 1985). The 4-merous apetalous flowers and opposite leaves are also unusual for the Rosaceae. Coleogyne ramosissima ______New growth and flowering are dependent on the quantity and distribution of precipitation. In the ab- Blackbrush sence of adequate fall or winter/spring rain, vegetative and reproductive growth are curtailed (Ackerman Blackbrush, the only member of its genus, is a long- and others 1980). Blackbrush enters a period of sum- lived densely branched shrub usually about 3 ft (0.9 m) mer inactivity in response to low soil moisture. Flower- in height, but sometimes reaching 6 ft (1.8 m) or more ing occurs from April to August depending on location. (fig. 7). The root system is shallow and spreading with Fruits ripen from April to June and are dispersed most roots less then 16 inches (40 cm) deep. Roots are shortly thereafter (Ackerman and others 1980). associated with the nitrogen fixing actinomycete Frankia (Bowns and West 1976; Dittberner and Olsen Ecological Relationships and Disribution 1983). The tangled ash-gray branches give the shrub a dark appearance, particularly when wet; hence the Considered a relict endemic species of arid and semi- name blackbrush (Benson and Darrow 1945). Main arid areas of western North America (Stebbins 1972), blackbrush’s present distribution is thought to represent a restriction of a once more extensive range (Stebbins and Major 1965). Blackbrush is distributed from southern California, east through southern Nevada, northern Arizona, southeastern Utah and southwestern Colorado in areas with annual precipitation in the 6 to 12 inch (150 to 300 mm) range (Harrington 1964; Munz and Keck 1959; Turner 1982; USDA Soil Conservation Service and USDI Bureau of Land Management 1981; Welsh and others 1987). Blackbrush occurs as a dominant species over extensive areas and is particularly common in the southwestern Great Basin Desert along the Colorado River drainage system (Turner 1982). Blackbrush is a major shrub along the Transition Zone between the Great Basin and Mojave Deserts (Tidestrom 1925). Monotypic stands within this Zone intergrade into mixed communities of Mojave Desert, Colorado Pla- teau, and Great Basin vegetation. Figure 7—Blackbrush community in south- Monotypic stands of blackbrush are characterized western Utah. by low understory species diversity and greater total

552 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs cover than most desert shrub communities. Plants are Seeds require a short wet prechilling period to re- long-lived and slow growing; their communities are lieve dormancy. Bowns and West (1976) reported highly stable. Blackbrush often replaces itself follow- prechilling at 39 ∞F (4 ∞C) for 23 days and incubation ing disturbances, but reestablishment of shrub cover at 57/40 ∞F (14/4 ∞C) provided 80 percent germination occurs very slowly due to extremely erratic seedling in 6 days and 90 percent in 13 days. Natural seedlings establishment and low growth rates (Webb and others are uncommon. Their appearance may be associated 1987). In southern Nevada, Beatley (1974) character- with dry autumns and winter or spring precipitation ized blackbrush as requiring Great Basin precipita- totaling 2 to 5 inches (50 to 130 mm) (Beatley 1974). tion and Mojave Desert temperatures and existing Plants can be established by artificial seeding, but primarily between creosotebush and pinyon-juniper sufficient plantings have not been attempted to ad- or sagebrush communities (Wells and Jorgensen equately determine appropriate seeding rates or seed- 1964). In Arizona, blackbrush occurs in areas having bed requirements. Local seed sources should be used; a mean annual soil temperature of 59 ∞F (15 ∞C) populations are not adapted to areas other than their (USDA Soil Conservation Service and USDI Bureau sites of origin. Seed can be planted with most conven- of Land Management 1981). tional seeders. A planting depth of about 0.5 inch In general, blackbrush stands occur on upland (1.3 cm) is recommended. Rodents may aid in natural sites, broad terraces, and gently rolling hills (USDA seed dispersal and planting, but rodent predation of Forest Service and USDI Bureau of Land Manage- planted seed has been observed (Bowns and West ment 1981). Soils are usually shallow, calcareous, and 1976). Fall planting is recommended to provide ad- coarse textured, ranging from gravelly to cobbly loam equate wet prechilling and reduce rodent problems or gravelly clay loam (Foster 1968; Kearney and Peebles (Bowns and West 1976). Stark (1966) reported good 1960; Shreve 1942). The shrub is less prevalent and success from seedings in Nevada, but wildland seedings occurs with an understory dominated by grasses in in Utah have not been as successful. Seedlings devel- drainage bottoms, where soils are moderately deep to oping from plantings in central Utah, north of the deep with fine sandy loam surface textures and calcar- species natural range, have failed to persist for more eous underlying layers of very gravelly sandy loam or than 2 to 3 years due to cold winter temperatures. gravelly clay loam (USDA Soil Conservation Service Seedlings developed slowly and some were suppressed and USDI Bureau of Land Management 1981). Other by herbaceous competition. In general, most attempts blackbrush sites with good fertility and soil water also to plant blackbrush offsite have not been successful. support a herbaceous understory (Bowns 1973). Only fair survival has resulted from planting Blackbrush normally occurs on soils low in salt con- projects using either container or bareroot stock. Con- tent (McCleary 1968). Seedlings are sensitive to salin- tainer stock grows moderately well in a buffered peat- ity, which may limit its distribution and spread vermiculite medium, but seedlings are sensitive to (Wallace and Romney 1972). overwatering. Four to 6 inch (10 to 15 cm) tall seed- Blackbrush is often a component of many transi- lings can be produced in 6 to 10 months under green- tional shrub communities that provide important for- house conditions. Survival has not exceeded 30 per- age and habitat for game and livestock. These include cent from plantings conducted in southeastern Utah. spiny hopsage, Nevada ephedra, rabbitbrush, big sage- Shading the small transplants improved survival brush, desert bitterbrush, cliffrose, and pinyon-juniper (Kitchell n.d.). communities. It is considered a potential candidate for use in revegetation; it occupies or once occupied exten- Uses and Management sive areas that have suffered severe abuse by destructive grazing practices. Dayton (1931) and Allen (1939) reported blackbrush provides fair winter forage for cattle, sheep, goats, Plant Culture deer, desert bighorn, and other game animals. Black- brush communities are commonly used as winter Blackbrush seed production is highly erratic. Most range for sheep. However, blackbrush is often re- stands produce a harvestable seed crop in perhaps garded as a poor forage species with low palatability 1 of every 6 to 10 years. Ripe seed readily detach from (Bates 1984; Sampson and Jesperson 1963). Although the plant and are dispersed by gravity. Seed collec- used by big game and livestock (Gullion 1964), it is tion is slow and tedious; consequently, commercial usually not a major component of their diet. It may supplies are rarely available. Fruits are cleaned become an important forage during severe winters using a fanning mill or gravity separator. Purity of (Kufeld and others 1973; Sampson and Jesperson 95 percent and germination of 70 percent are recom- 1963). Blackbrush is deficient in phosphorus for sheep mended for seed purchases. There are approximately and cattle and low in protein during fall and winter 27,015 seeds per lb (59,557 per kg) at 100 percent (Bowns and West 1976). In addition, the low growing, purity (see chapter 24). intricately branching habit of the shrub reduces its

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 553 Chapter 22 Rosaceous Shrubs attractiveness to livestock (Provenza and others 1983). (Bowns 1973; Bowns and West 1976). Most plants are Light to moderate grazing may reduce woody materi- killed if the understory is dense enough to carry a fire. als and spininess and increase production of palatable Low intensity burns may not kill mature plants. new growth (Provenza 1977; Provenza and others Bates (1984) reported considerable resprouting fol- 1983). Mule deer make moderate use of this shrub in lowing fires in California. Blackbrush seed may re- winter and spring (Leach 1956). Its seeds are used by main viable in the soil for several years; new seedlings birds and rodents (Mozingo 1987; USDA Forest Service may emerge from seed banks many years after mature 1937) and it provides cover for many small animals. plants are destroyed by fires or other disturbances Blackbrush is adapted to harsh sites, but its slow (Bates 1984). Blackbrush seedlings establish very growth and limited area of adaptation have restricted slowly following fires (Anderson 2001; Wright and its use in revegetation projects on non-native sites. All Bailey 1982); red brome and other annuals develop blackbrush sites do not respond similarly to restora- quickly and dominate burned sites, often precluding tion and management practices. Bates (1984) and blackbrush recovery (Rickard and Beatley 1965). How- West (1969) concluded that blackbrush occupies two ever, Bates (1984) observed blackbrush reestab- different situations, each responding differently to fire lishment from seed on sites occupied by annual and and other improvement techniques. In the first situa- perennial grasses following wildfires. Thus, the tion, blackbrush occurs in monotypic stands as a species may exert some competitiveness with herba- climax species supporting few native understory spe- ceous species in some locations. cies. Shrubs are widely spaced and cover provided by Jensen and others (1960) found burns in Nevada grasses and forbs may be extremely low. Cryptogamic effectively killed blackbrush plants. Unless native crusts covering interspaces reduce soil erosion; those perennial grasses and broadleaf herbs recovered rap- with a blue-green algae phycobiont provide an impor- idly following burns on these sites, annual weeds tant source of soil nitrogen (West and Skujins 1977). quickly dominated and prevented blackbrush recov- These sites are frequently considered for range im- ery. Some shrubs, including turpentine bush, desert provement projects. Burning has not been a successful bitterbrush, and desert peachbrush, are capable of means of releasing the existing understory due to the recovering following fires, and if present prior to burn- unpredictable recovery of native herbs, invasion of ing, may soon dominate burns where blackbrush was annual weeds, and loss of cryptogamic crust that once common (Bowns 1973). recovers very slowly (Jensen and others 1960). In the In California, annuals quickly invaded blackbrush absence of annual weeds, these sites often revert back burns, but were replaced by perennial herbs within to blackbrush without any intermediate stage (West 10 to 15 years (Bates 1984). A mixed shrub community and Skujins 1977). Attempts to plant substitute then appeared in about 30 years, eventually reverting shrubby species have not been successful. back to solid stands of blackbrush if the stands were Monotypic stands of blackbrush are often associated poorly managed. Not all sites responded so favorably. with shallow soils and soil water conditions that do Once annual weeds established, the reentry of more not favor seeding success. Annual weeds, principally desirable herbs and shrubs was often limited. Burning red brome, compete with the shrub seedlings. At- or chaining increased the density of other shrubs tempts to replant blackbrush back onto these sites present after treatment. following wildfires or other disturbances often meet Blackbrush communities provide desirable ground with failure. cover and soil protection and should not be removed or The second common revegetation situation occurs converted to other species unless the treatments are on sites where blackbrush invades other plant com- assured of some success. Anchor chaining has been munities or is naturally present in low density. Inva- effective in reducing blackbrush density where it ex- sion normally occurs on sites with deeper soils and ists in pure stands or intermixed with pinyon-juniper. better water availability than is found on monotypic Brush beating has also been used to decrease shrub blackbrush sites. Blackbrush increases on these sites density, remove older wood, and stimulate new growth when heavy grazing suppresses other plants. Under (Bowns and West 1976). Neither chaining nor brush these conditions, burning or chaining can be used to beating kills all shrubs. Double chaining for example, reduce blackbrush density and allow release of exist- initially killed about 25 percent of the plants on two ing natives or aid in the establishment of seeded large blackbrush chaining projects in southern Utah species. Success is dependent on edaphic and climatic (Monsen n.d.). Crested, intermediate, and pubescent conditions. Sites with deeper soils generally respond wheatgrass were seeded at both sites, producing good more favorably (Bates 1984; Halliday 1957). understory stands. Blackbrush resprouted and re- The successional patterns of recovery following fires gained its earlier productivity in about 10 years. Its are somewhat unpredictable in blackbrush communi- productivity was not suppressed by the presence of the ties. The shrub is reported by some to be a nonsprouter seeded grasses. Indian ricegrass has also responded

554 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs favorably following chaining of certain blackbrush black, two-celled pome about 0.4 inch (1 cm) long sites. Blackbrush is particularly compatible with In- (Johnson and Anderson 1980; Welsh and others 1987). dian ricegrass, and when protected from livestock grazing these sites often support a mixed grass-shrub Ecological Relationships and Distribution community (Jeffries and Klopatek 1987). Peking cotoneaster was originally introduced from Varieties and Ecotypes northern China in 1883 as an ornamental shrub (Hoag 1958; Leslie 1954; Slabaugh 1974). It has been Native stands in the Colorado River drainage in planted for conservation and shelterbelt purposes southeastern Utah provide the seed for most blackbrush throughout the United States, particularly in the revegetation projects. However, collections acquired northern Great Plains and upper Midwest. It is also from this area have not demonstrated adaptation to used in the southern Canadian prairie provinces other locations. If artificial plantings are contem- (Johnson and Anderson 1980; Slabaugh 1974). Like plated, seed should be acquired from native stands other Cotoneaster species, Peking cotoneaster is com- growing near the planting sites. Different ecotypes mercially cultured and sold for ornamental plantings. demonstrate specific attributes that appear to be ge- A number of varieties have been developed and are netic traits. Certain ecotypes demonstrate adaptabil- widely used. Plants have demonstrated a wide range ity to arid desert conditions; others express compat- of adaptation when planted in shelterbelts and conser- ibility with herbaceous understory plants, and some vation plantings in the West and Midwest. appear better able to recover from burning. Plant Culture Cotoneaster acutifolia ______Peking cotoneaster normally flowers in early May; fruits ripen in September or October (Slabaugh 1974). Peking Cotoneaster Plummer and others (1968) reported that fruits do not Peking cotoneaster is an unarmed, low-spreading ripen until mid November in Utah. Fruits remain on shrub, often reaching 12 ft (3.7 m) in diameter (fig. 8). the shrub until mid to late winter. Abundant crops are Plant height is usually less than 5 ft (1.5 m). The shrub often produced on cultivated hedges and shelterbelt is described as deciduous (Welsh and others 1987), but plantings. Fruits become rather dry as they ripen. numerous leaves often remain on the plant until late Mature fruits are easily dislodged from the shrub and winter. Branches are numerous, reddish brown, and are hand collected in late fall or early winter after arcuate to horizontal. Flowers are small, perfect, regu- some leaves have fallen. Seed are removed by mac- lar, and solitary or in cymes terminating lateral erating the fruits and flushing the pulp with water. branches. Flowers are white or pink. The fruit is a The remaining seeds and pulp are then dried and screened. Empty seeds are removed by flotation (Slabaugh 1974). Conditioned seedlots normally are very pure with little debris. The average number of cleaned seeds per lb varies from 21,984 to 32,310 (48,465 to 71,231 per kg) (Plummer and others 1968; Slabaugh 1974). Purity of 98 percent and germination of 80 percent are recommended for seed purchases. Germination of Peking cotoneaster is restricted by the presence of a hard, impermeable seedcoat and embryo dormancy (Slabaugh 1974). Scarification with sulfuric acid and wet prechilling pretreatments are recommended to hasten germination (Fordham 1962). However, fall nursery or wildland plantings usually provide sufficient wet prechilling. An alternating temperature regime of 50 ∞F (10 ∞C) for 15 hours and 77 ∞F (25 ∞C) for 9 hours, with light supplied during the warm cycle, usually provides optimal germination conditions for pretreated seed Figure 8—Peking cotoneaster, an introduction (Slabaugh 1974). from China, has received considerable use in Peking cotoneaster is commonly grown as nursery midwestern windbreaks and shelterbelts where stock for outplanting and is usually lifted after the it provides winter cover and food for birds and second growing season. Cuttings are easily propa- big game. gated; the plants layer readily. Wildings or rooted

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 555 Chapter 22 Rosaceous Shrubs stem layers can be harvested early in the spring for Varieties and Ecotypes field plantings. Doran (1957) reported softwood cuttings of cotoneaster taken in early summer root well. There are no releases. Wildland seedings have been only marginally successful. Seed germination is erratic. Seedlings grow Cowania stansburiana ______slowly and are easily suppressed by herbs. Planting success is improved if the seedbed is kept weed free. Stansbury Cliffrose or Cliffrose Established plants are more resilient and persist well. This shrub rarely suffers heavy damage from insects, Stansbury cliffrose has often been identified as a disease, or use by small mammals. Little damage has variety of C. mexicana var. stansburiana [Torr.] Jeps. been noted during heavy grasshopper infestations. However, we have elected to accept it as a separate species as proposed by McArthur and others (1983b). Uses and Management Plants are long-lived, erect, evergreen, aromatic shrubs or small trees 3 to 18 ft (1 to 5.4 m) tall with Peking cotoneaster has been most widely used for stiff, often somewhat brittle stems and gray shreddy windbreaks, shelterbelts, conservation, and ornamen- bark (Cline 1960; Welsh and others 1987). New growth tal plantings (Hoag 1958; Johnson and Anderson 1980; is flexible with glandular bark. Stansbury cliffrose has Leslie 1954; Slabaugh 1974; Wyman 1949). The shrub an extensive root system with long, spreading, and is used in multi-row windbreaks and cover plantings descending roots (Sutton and Johnson 1974). Roots to protect soils and reduce wind erosion. It furnishes a may form symbiotic nitrogen-fixing associations with low, dense groundcover that provides year-long pro- Frankia (Nelson and Schuttler 1984). Plants are res- tection. The plants are also commonly used in road- inous and strong smelling. The light to dark green way, recreation, or conservation plantings where leaves are clustered along the branches and are 0.5 to ground cover, low maintenance, and aesthetics are 1 inch (1.2 to 2.5 cm) in length with revolute margins important. Plants are attractive and provide glossy and five to seven teeth at the apex. Leaf surfaces are green foliage with persistent red or black berry-like glandular dotted and resinous above with a white pomes. The shrub withstands snow deposition and is tomentum beneath (Blauer and others 1975; Munz used to trap snow and protect roadways and buildings. and Keck 1959). About 90 percent of the showy, fra- Peking cotoneaster is commonly used to control erosion in residential areas, recreation sites, and wild- grant, cream to yellow flowers appear during the first lands. It is particularly useful in these areas as plants flush of flowering and are formed on lateral spurs of establish well from transplanting and quickly im- the previous year’s wood. Later flowers appear spo- prove ground cover. Transplants can be planted at radically throughout the summer near the tip of the various spacings depending on the availability of current year’s leaders (Shaw and Monsen 1983b). seedlings. Planted shrubs will spread to occupy the Later flowers are particularly abundant following sum- planting site, but do not seed into adjacent areas. mer rains. Flowers are about 0.8 inches (2 cm) in Peking cotoneaster is adaptable to many ranges diameter and are normally perfect with two series of where native species have been lost and are difficult to stamens enclosing 5 to 12 hairy pistils. Fruits are reestablish. It has been used to provide winter food for plumose achenes (fig. 9) dispersed by gravity and wind various birds and big game (Johnson and Anderson (Blauer and others 1975). 1980; Miller and others 1948). The plant exhibits high palatability to game and livestock and is moderately tolerant of grazing (Plummer and others 1968). It is readily grazed in midsummer by mule deer (Kufeld and others 1973; Leach 1956). Winter browsing has been heavy in Utah. Spring and fall use is light. Game animals are attracted to the shrub and can damage landscape plantings. Peking cotoneaster should not be used in roadway or other plantings where wildlife concentrations create hazards. Peking cotoneaster may be used to improve areas where game animals concentrate and more productive forage and cover is essential. This shrub can be planted to enhance depleted native communities and attract and reduce animal use of native species. It can also be planted to reduce erosion on disturbed sites and to protect watershed and soil values. Figure 9—Stansbury cliffrose branch with mature achenes, note the plumose styles.

556 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution concluded that two adaptive products have evolved through natural hybridization. Desert bitterbrush, Stansbury cliffrose is distributed from southern the southern adaptive derivative, is one product. The California east to Colorado and south to Sonora and second is less distinct. It occurs in areas north of Chihuahua (USDA Forest Service 1937; Welsh and Utah on sites not occupied by Stansbury cliffrose and others 1987). It does not occur north of about the 42 ∞N consists of antelope bitterbrush populations that ex- parallel (McArthur and others 1983b). It is found press Stansbury cliffrose characteristics, particularly primarily on dry, rocky, well-drained silty to gravelly low palatability. Stutz (1972) suggested selection pres- soils of foothills and mesas growing in blackbrush, sure resulting from heavy browsing has favored the chaparral, oak, desert shrub, mixed grass-desert shrub, less palatable plants. mountain brush, and pinyon-juniper woodlands up to Both Purshia and Cowania were present in the the edge of ponderosa pine forests at elevations from Madro-Tertiary geoflora, which evolved from the 3,000 to 9,000 ft (900 to 2,700 m) (Blauer and others Arcto-Tertiary and Neo-Tropical geofloras in response 1975; McArthur and others 1983b; Welsh and others to increasingly drier conditions in the Southwest 1987). (Axelrod 1958, McArthur and others 1983b). McArthur Cowania and Purshia are closely related genera but and others (1983b) postulated that the weak isolation differ in many physical characteristics (table 1) and barriers now existing between the two genera indicate geographical distribution (McArthur and others 1983b; that the present zone of contact of the two species Stutz and Thomas 1964; Thomas 1957). Because of developed after a long separation. Much of the area these differences, we prefer to retain the genus name where they now coexist was occupied by Pleistocene Cowania even though Welsh and others (1987) re- lakes and glaciers as recently as 12,000 years ago. cently transferred its species to Purshia. Disjunct flowering periods normally prevent hy- Plant Culture bridization between antelope bitterbrush and Stansbury cliffrose. Antelope bitterbrush usually flow- Although Stansbury cliffrose grows in semiarid re- ers earlier than Stansbury cliffrose (McArthur and gions it usually produces at least some seed each year. others 1983b; Shaw and Monsen 1983b), but aspect or Abundant or commercially harvestable seed crops site conditions may result in overlapping periods of develop in about 2 out of every 3 to 4 years. Although anthesis and pollination. Stutz and Thomas (1964) flowering usually occurs after the last spring frost, and Thomas (1957) reported that hybridization is so unusually late frosts can diminish seed yields. common that no populations of antelope bitterbrush The period of floral development, anthesis, and fruit in Utah have been examined that are free from the development lasts from 68 to 78 days in southern influence of Stansbury cliffrose. Introgression of ante- Idaho (Shaw and Monsen 1983b). Anthesis and polli- lope bitterbrush into Stansbury cliffrose is less com- nation occur over a 10-day period. Fruits then mature mon, but does occur. McArthur and others (1983b) about 37 days later. The plants are insect-pollinated and self-incompatible, consequently seed production is significantly impacted by high winds, rainy periods, unusually hot temperatures, and other factors that influence insect pollination. Table 1—Morphological characteristics separating Cowania Fruit (achene) ripening is less uniform for Stans- and Purshia (after Stutz and Thomas 1964). bury cliffrose than for antelope bitterbrush, but the Character Purshia Cowania bulk of the fruit can generally be harvested on one date. Fruits are hand collected by striking the limbs Twig pubescence heavy glabrous Dorsal leaf pubescence heavy glabrous with a paddle or stick to knock them into collection Hypanthium glands none 3+/mm2 hoppers or trays. The brittle styles are removed with Stalked glands absent present a Dybvig or barley debearder and separated by fan- Pistils 1 or 2 4+ ning. Seeds are not removed from the achene to avoid Style length <1.5 cm 1.6 - 3.4 cm damaging the embryo (Young and Evans 1983). Gen- Style pubescence puberulent plumose erally about one-third of the harvested material by L/W ratio of achenes <3.1 >3.3 weight consists of viable seed (achenes) (Alexander Stamen insertion hypanthium hypanthium and others 1974). There are about 64,600 seeds per lb margin throat (64,500 per kg) (Alexander and others 1974; Plummer Leaf margin revolution slight pronounced and others 1968). Purity of 95 percent and germina- Hypanthium pubescence pubescent glabrous tion of 85 percent are recommended for seed pur- Stamen series one two Lobes/leaf three five+ chases (see chapter 24). Leaf glandulosity none 3+/mm2 Seed can reportedly be warehouse stored for 5 to Leaf retention deciduous evergreen 7 years (Alexander and others 1974; Plummer and

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 557 Chapter 22 Rosaceous Shrubs others 1968; Springfield 1973), or even as long as 15 propagated from seed, but stem cuttings can be rooted years (Stevens and others 1981a). Germination tests (Shaw and Monsen 1983b). Because Stansbury cliffrose conducted on seed lots harvested from the same area is an evergreen shrub, plants are sensitive to lifting in different years exhibit similar germination pat- and transplanting if not properly hardened or dor- terns. However, some differences do occur. Poorer and mant. Survival of actively growing plants is generally slower germination results from unfavorable growing low. conditions during the year of seed production; more Grasshoppers utilize Stansbury cliffrose (Edgerton rapid and complete germination characterize seed and others 1983) but have not been observed to seri- produced during years of abundant production ously damage or eliminate new plantings. Weedy (Jorgensen n.d.). Shaw and Monsen (1983b) reported competition on the planting site should be controlled the loss of achenes prior to the time of maturation by burning, mechanical, or other means of seedbed was 13 percent in 1979 and 52 percent in 1980. preparation prior to transplanting (Giunta and others Although insects were a major factor, losses could not 1978; Shaw and Monsen 1983b). be attributed to insect damage alone. Stansbury cliffrose seed requires a wet prechilling Uses and Management treatment to release dormancy. The optimal treatment duration varies with population and year of Stansbury cliffrose is regarded as a desirable winter harvest. Consequently, seed should be planted in fall forage for big game as well as cattle and sheep (Blauer or winter; spring seeding should be avoided. Meyer and others 1975; Plummer and others 1968; Robertson and Kitchen (n.d.) found that the rate and total germi- 1974; Tueller 1979; USDA Forest Service 1937). Kufeld nation percentage increased from 30 percent after 2 and others (1973) reported that mule deer make heavy use of it in fall, winter, and spring but only moderate weeks of wet prechilling at 33 ∞F (1 ∞C) to 100 percent in 7 days after 6 weeks of wet prechilling. use in summer. Some Stansbury cliffrose stands are grazed by big game and livestock during all seasons. Seed maintained at 33 ∞F (1 ∞C), began germinating during prechilling within 8 weeks and completed However, the species is not as universally palatable as germination in 11 weeks. Plummer and others (1970b) antelope bitterbrush (McArthur and others 1983b; reported 89 percent germination after 90 days at Stutz and Thomas 1964). Plants often receive negligible use in spring and summer in areas where other alternating temperatures of 32/85 ∞F (0/31 ∞C) and 99 percent germination after 90 days of alternating forage is available (USDA Forest Service 1937). Cer- tain ecotypes, including some southern Arizona popu- temperatures of 32/38 ∞F (0/3 ∞C). Young and Evans lations, evidently have low palatability in all seasons. (1981b) recommended prechilling at 32, 36, or 39 ∞F (0, Two dimensional chromatograms of their phenolic 2, 4 ∞C) for approximately 6 weeks. Seeds do not require light for germination (Sabo and others 1979). compounds can be distinguished from those of more Stansbury cliffrose is generally fall seeded in a palatable populations (Blauer and others 1975). mixture with other shrub seed. Seed can be planted Like other evergreen species, Stansbury cliffrose with most conventional drills or seeders. Cleaned maintains its protein levels and energy values in seed must be incorporated into the soil to a depth of winter (Dittberner and Olsen 1983; Smith and Hubbard about 0.25 to 0.50 inch (6 to 12 mm) (Alexander and 1954; Welch and others 1983a). In a common garden others 1974; Plummer and others 1968). Surface broad- study in southern Idaho, Stansbury cliffrose leaders casting is generally unsuccessful if the seed is not were found to contain a significantly higher percent- subsequently covered. Seedling emergence is similar age of crude protein in winter (8.4 to 8.6 percent, dry to that of antelope bitterbrush, occurring early in weight basis) than those of antelope bitterbrush or spring. Small seedlings are sometimes subjected to Apache plume, but slightly less than those of desert late frosts. During the establishment period, seed- bitterbrush (Welch and others 1983a). The in vitro lings are quite sensitive to competition and water digestible dry matter of Stansbury cliffrose exceeded stress. However, once established, Stansbury cliffrose that of all species tested (37.6 percent). seedlings tolerate drought, heat, and soil salinity Stansbury cliffrose can withstand heavy grazing better than antelope bitterbrush or Apache plume and does occur in areas where winter use is quite (Plummer and others 1968). Seedlings are not capable intense. Flowering and production of new growth may of competing with cheatgrass, red brome, or dense be stimulated by 65 percent use; browsing in excess of stands of some perennial grasses. 80 percent of new growth decreases vigor (Wasser Stansbury cliffrose is easily grown as bareroot (Shaw 1982). Stands frequently sustain game animals dur- 1984) or container stock (Landis and Simonich 1984); ing years of heavy snow accumulation and should be it develops rapidly if properly cultured. One-year-old managed to ensure forage availability during the bareroot seedlings generally attain satisfactory size critical midwinter period. for field transplanting. Container plants are usually

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Stansbury cliffrose has been seeded on a variety of Seedlings of Stansbury cliffrose must be protected sites to improve diversity and winter wildlife habitat from excessive competition and heavy grazing until value of vegetation (Edgerton and others 1983). Good plants are 3 to 5 years of age. In addition, Stansbury stands can be established in the mountain brush zone cliffrose stands have been and are being lost due to the and in most pinyon-juniper communities (Davis 1983; invasion of weedy annuals that restrict natural or Giunta and others 1978; Monsen and Christensen seeded shrub establishment and expansion and ulti- 1975). Stansbury cliffrose is frequently intermixed mately contribute to the loss of the shrub overstory with Wyoming big sagebrush, blackbrush, and salt (Cline 1960; Price 1982). Sites should be managed or desert shrubs and can be successfully reseeded on treated to create a weed-free seedbed and maintain a such sites. Although Stansbury cliffrose frequently suitable native understory after planting. grows on soils derived from limestone, it is not limited Stansbury cliffrose frequently does not recover from to such sites. Establishment and survival are less burning, chaining, or other methods of mechanical certain when the species is planted in shrub commu- treatment. Multiple stemmed shrubs are more toler- nities where it does not occur naturally. However, ant than older single-stemmed shrubs (Plummer and plantings have established and survived successfully others 1968). Extreme variability in response to fire for nearly 40 years in antelope bitterbrush communi- has been reported (Ferguson 1983; Howard 1995; ties in south central Idaho and for over 10 years in Wright and others 1979), perhaps reflecting differ- northeastern Oregon and eastern Washington (Blauer ences among ecotypes. Attempts to improve existing and others 1975; Edgerton and others 1983; Monsen stands using these treatments have not been success- and Shaw 1983c). Stansbury cliffrose is not fire toler- ful unless the sites are also seeded. ant and should not be planted in areas subject to repeated burning. Varieties and Ecotypes On native sites Stansbury cliffrose may function as a pioneer species following disturbances (Wright and The extensive natural hybridization of Stansbury others 1979). It has been considered a candidate plant cliffrose with antelope and desert bitterbrush, and to for mined areas, roadways, and other disturbed sites. a lesser extent Apache plume, provides an immense It provides only moderate erosion control due to its array of material for improving palatability; increas- upright habit and low growth rate (Dittberner and ing drought, heat, and fire tolerance; broadening areas Olsen 1983; Smith and others 1978). Its frequency of of adaptation; enhancing evergreen growth character- nodulation by the nitrogen-fixing endophyte is gener- istics and forage traits; and manipulating growth ally low and varies with collection date and location habits of these taxa (McArthur and others 1983b; (Nelson 1983). Plants have been successfully estab- Monsen and Davis 1985; Stutz 1972). Improvement of lished on mine sites near Alton, UT, and on the seedling vigor, establishment, and initial growth rates phosphate mines of southeastern Idaho (Ferguson and would enhance the use of Stansbury cliffrose and its Frischknecht 1985; Monsen, data on file, Shrub Science hybrids. Lab., Provo, UT). Replacement of topsoil is required to Stansbury cliffrose selections from central Utah attain satisfactory stands. Results of roadway seedings have been among the most successful and productive and transplanting trials in Utah and Idaho have been populations seeded on pinyon-juniper sites in Utah erratic, even when plantings are conducted in areas (Monsen and Davis 1985). Hybrids with antelope where Stansbury cliffrose naturally occurs. bitterbrush collected from Utah County, UT, also Stansbury cliffrose is an attractive, drought-tolerant offer excellent forage traits, including evergreen growth, high productivity, dramatic regrowth, good ornamental. The leaves are evergreen, the flowers and palatability, and an upright growth form. Davis fruits are showy, and flowering occurs over a pro- (1983a) listed five different central Utah accessions longed period. Mature shrubs can become rather ragged of Stansbury cliffrose that have performed best in in profile, but respond well to pruning. artificial seedings in Utah nearly 30 years after Stansbury cliffrose seedlings grow slowly and may planting. Studies were based on continuous evalua- require 3 to 4 years to reach a height of 6 to 12 inches tions from 60 sites and include such factors as growth, (15 to 30 cm) (Monsen and Christiansen 1975). Growth persistence, and vigor. thereafter is much more rapid. In a big game exclosure Stansbury cliffrose introgressions with desert bit- near Boise, ID, Stansbury cliffrose planting stock terbrush acquired from Benton, CA, demonstrate a grew more slowly than several sources of antelope wide range of adaptation and survival under harsh bitterbrush for the first 4 to 5 years, but exceeded circumstances. Selections from hybrids with desert their growth rates thereafter. Both species produced bitterbrush show promise for improved fire tolerance seed 5 to 6 years after planting (Shaw and Monsen and adaptation to arid situations. 1983b).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 559 Chapter 22 Rosaceous Shrubs

Crataegus douglasii ______of the subfamily Pomoideae, their chromosome number is n = 17 (McArthur and Sanderson 1985). Douglas or River Hawthorn Douglas hawthorn, the most common species in our area, is distributed from Alaska east to Alberta and Douglas or river hawthorn is a compact, deciduous, the Dakotas and south to California and northern deep rooted shrub or small tree growing 3.3 to 13 Wyoming. It also occurs in Michigan and Ontario (26) ft (1 to 4 [8] m) tall (fig. 10). The bark is smooth (Hitchcock and others 1961). In sagebrush and ponde- and reddish brown, becoming reddish bronze in win- rosa pine communities it is largely confined to riparian ter. Stems are armed with stout, usually straight zones where it may form dense thickets. It is well thorns 0.4 to 0.8 inch (1 to 2 cm) long. The petiolate adapted to deep, moist, organic soils, but may be found leaves are 1.2 to 2.4 inches (3 to 6 cm) long, usually less on soils ranging from silty to rocky clay loams. Al- than half as wide, and lanceolate to elliptic ovate. Leaf though it commonly grows on well-drained sites, Dou- edges are serrate to doubly serrate. Leaves turn bright glas hawthorn is capable of withstanding periods of red in fall. Flowers are perfect, regular, and epigy- soil saturation (Stark 1966). The species is an alter- nous, developing in corymbose cymes that terminate nate host for cedar apple rust and may be damaged by short lateral spurs. The clusters of fragrant white leaf spot in drier locations (Krebill 1972; Stark 1966). blossoms sometimes appear to cover the crown of the plant. Fruits are lustrous black, glabrous pomes about Plant Culture 0.25 inch (0.6 cm) in diameter that resemble small apples. They are fleshy, dry, and mealy, and contain Ripe fruits are hand collected. Freshly collected light yellow pulp with one to five seeds. Flowering fruits must be kept cool and not allowed to mold prior occurs in May and June; fruits ripen from July to to extraction. Seed can be separated from fleshy mate- September (Brinkman 1974e; Hitchcock and others rial with a Dybvig cleaner or blender and water fol- 1961; Welsh and others 1987). Fruits are dispersed by lowed by air drying and fanning. A variable percent- gravity and animals. age of the seed is usually unsound but cannot be removed by flotation. Some of these may be separated Ecological Relationships and Distribution on a gravity table. There are about 15,050 seed per lb (33,179 per kg) at 100 percent purity (see chapter 24). There are approximately 300 species of Crataegus Treatment with concentrated sulfuric acid has been distributed throughout the Northern Hemisphere. In used to release dormancy. The appropriate length of North America the genus is most abundant east of the acid treatment varies from 0.5 to 3 hours depending on Rocky Mountains. However, approximately 8 to 10 the seedlot (King 1947; McKeever 1938). Effective- species occur in the Rocky Mountain area (Preston ness of acid scarification has not been consistent; acid 1968). Hawthorn taxonomy is extremely complex; the genus consists of numerous species exhibiting great can penetrate the seed coats and damage the embryos variation. Hybridization is common, particularly in of fresh seed. Embryo dormancy is released by a 3 to the central and eastern United States. Characteristic 5 month wet prechilling (King 1947; McKeever 1938; SEAM 1976). Because of the variability in dormancy among seedlots, combinations of standard pretreatments may be only partially effective in its release. Tetrazolium or excised embryo tests are often used as indicators of seed quality because current germination tests are time consuming and not highly reliable. Results of direct seedings have been erratic. Spot plantings in desirable locations conserve seed and provide developing seedlings with the best opportunity for establishment. Douglas hawthorn is most reliably established from planting stock. Container and bareroot seedlings are easily propagated, handled, and planted with good survival on adapted sites. In the bareroot nursery, seed should be planted to provide a seedling density of 25 per ft2 (278 per m2). Untreated seed should be planted in early fall, but wet prechilled seed must be spring planted (Hartmann and others 1990; Van Dersal 1938). Container seedlings of uniform size are most Figure 10—Douglas hawthorn is an excellent economically produced from germinants (Landis and wildlife species that often occurs in or near Simonich 1984). Rooting of hardwood or softwood riparian areas.

560 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs cuttings is not recommended (Marchant and Sherlock It is distributed from British Columbia and Alberta 1984). Root cuttings have been used to propagate south to Oregon, Idaho, and Montana. It grows well in cultivated ornamental Crataegus species (Wyman heavy clay soils over basalt. Yellow hawthorn is found 1971). Seedlings may be planted in mixtures with from central Utah east to Pennsylvania and south- other shrubs, but should be protected from competi- eastern Canada. tion with perennial grasses and weedy species.

Uses and Management Fallugia paradoxa ______Douglas hawthorn is a valuable shrub deserving Apache Plume more frequent inclusion in revegetation projects. It Apache plume, the only species of its genus, is a has been successfully used in conservation and wild- much-branched shrub 2 to 7 ft (0.6 to 2.2 m) tall life projects to furnish soil protection and food and (fig. 11). Branches are white to straw colored with cover for wildlife. It is well suited to mixed plantings flaking or scaly bark (Harrington 1964). Leaves are with other shrub species. The dense thickets provide described as often evergreen in the southern portion of hiding and thermal cover food for birds, small mam- its range (Blauer and others 1975; Deitschman and mals, bear, big game, and livestock. They add valuable others 1974b; USDA Forest Service 1937) and persis- structural diversity on grassland sites. Winter mule tent or deciduous in cooler areas (Munz and Keck deer use was evaluated as moderate to heavy in 1959; Welsh and others 1987). Leaves are borne in Washington (Brown and Martinsen 1959). Douglas fascicles along the branches and are 1.2 to 1.8 inches hawthorn receives moderate summer use by mule (3 to 5 cm) long. They are cuneate with revolute deer (Kufeld and others 1973). Use by white-tailed margins and pinnately divided into three to seven deer and livestock is variable (Van Dersal 1938). linear lobes (Welsh and others 1987). Leaves are Douglas hawthorn is rated as a valuable honey plant lepidote on both surfaces, light green above and rusty and fruits are heavily used by birds. below (Harrington 1964; Munz and Keck 1959; Welsh Fruits of Douglas hawthorn are high in sugar, but and others 1987). The large showy, perigynous flow- low in fats and protein. They were eaten fresh or dried ers, 1.2 to 1.8 inches (3 to 4 cm) broad, develop singly and used in pemmican by Native Americans. They are or in few-flowered cymes. A typical flower has five sometimes used for jams and jellies (Craighead and broad sepals alternating with five bracts and five others 1963). Where adapted, Douglas hawthorn is an large white petals. Numerous stamens are inserted in excellent candidate for landscape plantings in parks three series on the margin of the hypanthium. There and recreation areas. are 20 to 30 pistils. The hairy achenes are 0.25 inch (6 Natural reproduction from seed is often prevented mm) long with plumose persistent styles that elon- by heavy use and trampling in riparian areas where gate to 1 to 2 inches (2 to 5 cm) at maturity (Deitschman game and livestock tend to congregate. Seedlings and others 1974b; Young and Young 1986). The dis- develop slowly and should be protected from browsing tinct plumose styles resemble an Indian war bonnet, and competition with weeds and grasses during the hence the name “Apache plume.” first two seasons. Young seedlings may be girdled by rodents. Established plants are not easily damaged by excessive browsing.

Varieties and Ecotypes Although Douglas hawthorn exhibits many good attributes, selection trials have been limited. Appro- priate growth forms of Douglas hawthorn may be selected for shelterbelt and windbreak plantings. The deep rooted, thicket-forming plants are useful for stabilizing mine spoil banks, riparian sites, and other disturbances. They are resistant to beaver damage (Marchant and Sherlock 1984). A number of hawthorn species and varieties are native to riparian zones in the Intermountain region and may be selected for the revegetation of adapted sites (Hitchcock and others 1961). Columbia hawthorn is a small tree or shrub Figure 11—Apache plume is a resprouting rela- with red fruit and thorns 1.5 to 3 inches (4 to 7 cm) long. tive of bitterbrush that often grows in dry washes.

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Flowers are bisexual or rarely staminate (McMinn 1960). Plants established in southwestern Idaho flow- 1951). Blauer and others (1975) found monoecious as ered erratically from May until early October (Shaw well as dioecious populations. At a study site near and Monsen 1983b). Time from the appearance of Richfield, UT, Blauer and others (1975) encountered floral buds to fruit ripening was 67 days in 1979 and 86 shrubs with mostly perfect flowers. However, some days in 1980. plants had well-developed stamens but rudimentary Pollination is insect dependent; the flowers attract a and nonfunctional pistils. Other plants bore flowers variety of colorful bees, ants, hornets, and beetles with well-developed pistils, but rudimentary and non- (Blauer and others 1975). Good seed (achene) crops functional stamens. Plants grown from seed collected occur at 1 to 3 year intervals (Deitschman and others at this location exhibited similar differences when 1974b). Because flowers of Apache plume appear in grown at a study location near Boise, ID (Shaw and late spring, seed crops are usually not damaged by Monsen 1983b). McArthur (n.d.) considers many spring frosts. Determining the appropriate date to populations to be functionally dioecious. harvest seed is difficult; seeds ripen unevenly over a period of several months. Most flowers develop early Ecological Relationships and Distribution in the season, producing up to 90 percent of the annual seed crop. As seeds ripen they are detached and dis- Apache plume is found on a wide variety of sites from persed by the wind. Consequently, the period for southeastern California to southwestern Colorado collecting the maximum amount of seed may be rather and south into Mexico and western Texas, largely short, perhaps only 1 to 2 weeks in duration. Later outside the Great Basin (Deitschman and others blooming is associated with summer rains. 1974b). It is more abundant in the southern portion of Achenes turn from green to reddish as they ripen and plumes sometimes change from reddish to white. its range. Apache plume grows in a variety of plant They are harvested by dislodging the fruits from the communities from the lower desert brush types up bush. The entire fruit, including the achene and plume through the pinyon-juniper woodlands to open ponde- is collected. Fruits comprise only 15 to 20 percent of rosa pine communities at elevations from 4,000 to the collected material by weight. Unless the plumes 8,500 ft (1,200 to 2,550 m) (Dietschman and others are removed with a barley debearder, the collected 1974b; Monsen and Davis 1985). It is commonly found material remains in a thick entangled mass that in washes and ephemeral waterways, but also occurs cannot be seeded. There are approximately 546,000 on rocky or gravelly slopes and alluvial fans. Plants seeds (achenes) per lb (1,203,712 per kg) at 100 per- tend to be widely spaced, even when in pure stands. cent purity. For seed purchases, acceptable purity is 80 Apache plume superficially resembles Stansbury percent and germination is 75 percent (see chapter 24). cliffrose; both have lobed leaves and clusters of long Apache plume has been somewhat difficult to estab- plumed fruits. Unlike Stansbury cliffrose, Apache lish from direct seeding. A better understanding of the plume produces functionally dioecious plants (Blauer species’ seed germination requirements and seedbed and others 1975; McArthur and others 1983b), and it ecology would facilitate its use. Seeds are nondormant does not form symbiotic nitrogen fixing root nodules and germinate quite readily. Plummer and others with the actinomycete Frankia spp. (Klemmendson (1969) reported 60 and 70 percent germination after 1979; Nelson 1983; Righetti and others 1983). The 60 days at temperatures of 32 and 38 ∞F (0 to 3 ∞C). species occur in broadly overlapping areas (Benson Deitschman and others (1974b) found seeds sown in 1957; Blauer and others 1975), and a few suspected nursery beds germinated within 4 to 10 days. natural hybrids have been reported (Blauer and Although the plumed seeds are wind dispersed and others 1975). Some seeds have been produced by adapted to shallow or surface seeding, broadcast seed- artificial crosses, but no progeny have been produced ing without seed coverage has not been a satisfactory (Blauer and others 1975). Chromosome number for means of planting. Apache plume is generally in- Apache plume is n = 7 compared to n = 9 for Stansbury cluded in shrub seed mixtures that are planted sepa- cliffrose. rately from grasses. The seeds are easily sown with conventional drills if the styles are removed. Seed Plant Culture should be planted 0.25 to 0.5 inch (6 to 12 mm) deep in a firm seedbed. Fall seeding or seeding prior to the Flower buds are formed on elongated peduncles season of most dependable soil water is recommended. produced from vegetative growth of the current sea- Rodents often gather and bury or eat seeds that have son. Buds may appear on terminal or lateral leaders or fallen from shrubs as well as sown seed. Seeding in late from new root sprouts (Rehder 1940). Flowering be- fall or winter when rodents are less active reduces gins as early as April but may extend into August these losses. (Rydberg 1954; Van Dersal 1938). Plants flower from Seedlings emerge and grow rapidly under favorable April to June in California (McMinn 1951), and as conditions. They are reasonably drought tolerant, late as October in Arizona (Kearney and Peebles especially those that persist through the first summer.

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The small plants are not able to persist amid herba- by livestock on most sites. However, because of its fine ceous competition. twigs and evergreen leaves, it is a fairly valuable winter Bareroot and container reared seedlings are easily forage, particularly in the southeastern portion of its started and grow rapidly with irrigation, often flower- range (Ferguson 1983; Monsen and Davis 1985; USDA ing during the first growing season. When grown Forest Service 1937). The bushy habit provides good under field or nursery conditions in northern climates, cover for birds and mammals (Blauer and others 1975). the seedlings are mostly deciduous. Once the leaves Ferguson and Frischknecht (1985) considered Apache have fallen, the plants are dormant and can be lifted plume a secondary species for use on mine distur- and transplanted. The brittle stems must be handled bances in northeastern Utah. It is not nodulated by the carefully to prevent damage. In addition, the root nitrogen-fixing endophyte Frankia (Nelson 1983; system usually consists of a main root with few side Nelson and Schuttler 1984). Because of its ability to branches. Any damage to the roots can be detrimental spread vegetatively, Apache plume can be used to to field survival. Greenhouse-maintained plants tend provide ground cover and soil protection in dry regions to remain evergreen. These must be adequately hard- where infrequent storms can cause considerable ero- ened or overwintered in a shade house and planted as sion (Thornburg 1982). It is capable of recovering by dormant stock or field survival will be reduced. vigorous crown sprouting. On native sites it spreads naturally onto disturbed areas. Apache plume provides diversity for landscape Uses and Management plantings, particularly for dry, low maintenance situ- Apache plume occurs throughout the dry, hot por- ations. The white branches and the flowers and plumes tions of the pinyon-juniper, big sagebrush, and south- are showy and quite unusual. Under cultivation the ern desert shrublands. It is a useful species for these plant can be over irrigated and will not withstand difficult-to-plant areas and can be planted on southern even partial shade. Increased sprouting and a thicker, aspects and shallow soils. To date, Apache plume has bushy habit can be obtained for landscape or erosion most frequently been planted on drier portions of control plantings by trimming or grazing the planting. pinyon-juniper communities in central and southern Apache plume has been cultivated as far north as Utah. Seed of the more northerly-occurring ecotypes Massachusetts (Deitschman and others 1974b). is harvested for these projects. Once established The Tewa Indians of New Mexico used bundles of the Apache plume also does well on sites within its range smaller branches of Apache plume as brooms and the occupied by green ephedra, Stansbury cliffrose, and larger straight branches as arrow shafts (USDA For- black sagebrush. It can be planted on benchland sites est Service 1937). where spiny hopsage, shadscale, blackbrush, low rabbitbrush, desert bitterbrush, littleleaf mountain ma- Varieties and Ecotypes hogany, and singleleaf ash may occur. It is also adapted to fourwing saltbush, winterfat, and roundleaf Ecotypic differences have been observed among buffaloberry sites. The species has been established Apache plume collections currently under test. Shrubs of different populations vary in size and growth habit. from seed or planting stock on big sagebrush and antelope bitterbrush/bunchgrass ranges in central Some populations appear better adapted to more north- and northern Utah, southwestern Idaho (Shaw and erly climates than others. Breeding system expression, palatability, and rootsprouting following fire also Monsen 1983b), and northeastern Oregon (Edgerton and others 1983), all outside its native range. vary. There are at present no released varieties of the Apache plume does not form thick stands. When species. seeded with other shrubs it quickly thins out to occupy only specific microsites. This characteristic has also Holodiscus discolor ______been observed when it is planted on mine wastes and road disturbances. Apache plume has been slow to Creambush Oceanspray spread from seed, particularly when planted outside its natural range. Few seedlings are noted, even fol- Creambush oceanspray, also known as arrowwood, lowing years of heavy seed production. On coarse hardhack, rockspirea, and mountain spray, is a highly soils, vegetative spread occurs through root prolifera- variable, but usually erect multiple-stemmed, decidu- tion and root sprouting. Sprouts are more numerous ous shrub (fig. 12). Mature plants range from 2.5 to during years with good precipitation or following 10 (20) ft (1 to 3 [6] m) in height, with large tree-like physical damage to the plant. Individual sprouts may forms growing in coastal areas. Young twigs are fine attain heights of 1 to 2 ft (30 to 70 cm) in one season hairy; bark of the older stems is gray to deep grayish (Monsen n.d.). red. Leaf buds are covered by two to three purplish- Although Apache plume’s palatability to cattle and brown bud scales. The alternate leaves are 1 to 4 sheep is generally considered low, it is rated as good on inches (2.5 to 10 cm) long and ovate to ovate lanceolate some ranges. It does not receive heavy summer browsing with lobed to doubly toothed edges. Upper surfaces are

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rocky talus slopes and along streams in drier areas (Patterson and others 1985). Creambush oceanspray is associated with drier conifer types of the interior and is a climax species in a number of forested communities (Cholewa and Johnson 1983; Cooper and others 1987; Halverson and others 1986; Steele and others 1981). An understory shrub layer of mallowleaf ninebark and creambush oceanspray is common in ponderosa pine and Douglas-fir forests in eastern Washington, northeastern Oregon, northern Idaho, and western Montana (Daubenmire and Daubenmire 1968; Steele and others 1981). Creambush oceanspray occurs as a climax in open stands of conifers; it survives fires and other disturbances by resprouting from buds on the root crowns to become a member of early seral brushlands (Mueggler Figure 12—Creambush oceanspray in full bloom. 1966; Stickney 1986). Mixed seral brushfields provide valuable forage and cover for deer, elk, and other wildlife. Regeneration from seed following burns is often limited and seedlings develop slowly (Ferguson 1983; Wright and others 1979; Young 1983). However, hirsute, while lower surfaces are pilose lanate to regeneration from seed may be important following lanate and sometimes glandular. The common name extremely hot fires (Stickney 1974a). Owens (1982) creambush oceanspray is derived from the appear- found weight and length of current annual leader ance of the diffuse, showy panicles of tiny, perfect, growth increased directly with degree of canopy mor- perigynous, white to pink flowers. Panicles are 4 to 7 tality. However, regrowth was slightly lower for com- (12) inches (10 to 17 [30] cm) long and overtop the plete canopy mortality than for the 51 to 99 percent shrub. Flowers are about 0.2 inch (5 mm) in diameter, mortality class. Prescribed burning at 10 to 15 year regular, and 5-merous; petals exceed the sepals. Dried intervals has been recommended to rejuvenate brush flower parts and panicle branches turn brown and fields that have grown beyond the reach of browsing persist well into the winter. Fruits are light yellow animals (Orme and Leege 1980). hirsute achenes approximately 0.1 inch (2 mm) long. In communities where it exists, creambush Plant Culture oceanspray is one of the first shrub species to leaf out in spring (Orme and Leege 1980). Floral buds develop Collection of creambush oceanspray achenes is te- in late spring and flowering occurs from late June to dious; supplies are rare and costly. Fruits are “combed” late August, depending on location. Fruits mature in from the branches by hand in late fall and conditioned late summer and may persist into fall. Seeds are wind by air drying and screening out larger material with a dispersed (Hitchcock and others 1961; Orme and Leege fanning mill. Achenes of oceanspray are among the 1980; Stickney 1974a; USDA Forest Service 1937). smallest of shrub fruits. Only a small percentage of the seeds are normally sound; these are identified by Ecological Relationships and Distribution examining wet seed through a dissecting microscope for the presence of an embryo (King 1947). Seeds The genus Holodiscus existed in the Madro-Tertiary require wet prechilling at 34 to 36 ∞F (1 to 2 ∞C) for geoflora of western North America. It is now dis- about 120 days to break dormancy (Marchant and junct across the tropics, and occurs in western North Sherlock 1984; Young and Young 1986). America and Columbia. Creambush oceanspray is Seeding may be accomplished by broadcasting fresh distributed from British Columbia east to western achenes over a rough seedbed in fall and allowing Montana and south along the coast to southern them to be covered by natural soil sloughing. Achenes California. It grows in northeastern Oregon, Idaho, may be mixed with seeds of other shrub species, but and northeastern Nevada (Hitchcock and others 1961; they should not be sown with grasses or forbs. Spots to Munz and Keck 1959; Stark 1966; USDA Forest Ser- be planted should be carefully selected to make best vice 1937). Creambush oceanspray grows on a wide use of the seed. Seeding results have been erratic. variety of sites at elevations ranging from sea level to Creambush oceanspray is readily propagated from 7,000 ft (2,100 m). It is adapted to rich, deep soils of bareroot or container stock. Achenes should be fall coastal forests, riparian areas, and canyon bottoms. It sown in the nursery or artificially prechilled and may also be found growing on dry, shallow soils or spring sown. Cleaned achenes can be planted at

564 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs reasonably uniform spacings. They can be surface for one or two growing seasons. Excessive browsing is sown and covered by dragging a lightweight chain over rarely a threat to developing seedlings, but they may the seedbed. Seedlings develop rather slowly and may be girdled by rodents or trampled by big game or be lifted as 1-0 or 2-0 stock depending on size specifi- livestock. cations and growing conditions. Container seedlings are propagated by planting germinants or about 10 Varieties and Ecotypes wet prechilled seeds in each container. Plants can be grown from cuttings, but techniques have not been To date, selection work with Holodiscus species has been limited. The erosion control and ornamental refined (Everett and others 1978a). Marchant and potential of both creambush oceanspray and rockspirea Sherlock (1984) achieved 19 percent rooting success deserve further investigation. with softwood cuttings harvested in late June and propagated under a greenhouse mist system and 12 percent rooting success with hardwood cuttings over- Holodiscus dumosus ______wintered in a cold frame. Rockspirea Uses and Management Rockspirea, bush oceanspray, or rock spray spiraea is Creambush oceanspray is generally not considered somewhat similar in appearance to creambush ocean- a palatable shrub, particularly when alternative browse spray, but has a lower and more compact growth habit species are available (Daubenmire and Daubenmire (fig. 13). The rounded plants branch from the base and 1968; USDA Forest Service 1937). It is taken by deer range from 3 to 12 ft (1 to 4 m) in height with crown and elk; peak use occurs in fall and winter (Stickney diameters of 5 to 10 ft (1.5 to 3 m) (Hitchcock and 1974a). Use varies geographically depending on ecotype others 1961; Sutton and Johnson 1974; Welsh and and associated species. Creambush oceanspray re- others 1987). Inflorescences and leaves are smaller. ceives year round use by elk and deer in eastern Leaves are decurrent along the petioles; blades are Washington, where it forms an important component shallowly or coarsely toothed, sometimes with a few of the available browse (USDA Forest Service 1937). secondary mucronulate serrulations. Roots are densely Garrison (1953) recommended that browsing be lim- fibrous and spreading. Flowering occurs unevenly all ited to 50 to 60 percent of current annual growth on summer (Mozingo 1987). The species has n = 18 these sites. In the northern Rocky Mountains, use by chromosomes (McArthur and Sanderson 1985). big game is limited, but the species can be important Rockspirea is distributed from northeastern on some winter ranges (Leege 1969; Stickney 1974a). California across central and southern Oregon and Use increases tremendously the first winter following Idaho to Wyoming and south to Chihuahua, Mexico fires, but decreases the second winter (Leege 1969). (Hitchcock and others 1961) at elevations from 2,500 Palatability of creambush oceanspray is rated fair for to 12,000 ft (762 to 3,660 m) (Dittberner and Olsen cattle and poor to fair for sheep (USDA Forest Service 1983; Harrington 1964; Hopkins and Kovalchek 1983). 1937). The species tends to increase on heavily grazed It occurs in a wide array of plant communities including sites (Hall 1973; USDA Forest Service 1937). The quantity of the plant consumed in summer and fall in these areas may be considerable due to its abundance and accessibility. Creambush oceanspray is a useful erosion control plant. The species is adapted to steep, dry, rocky areas with shallow, sandy to clay loam soils (Stark 1966). Seedlings quickly develop a spreading, highly branched fibrous root system. Their low palatability and their ability to resprout following top damage contribute to their value in stabilizing critical sites such as roadcuts and fills (Hungerford 1984; Stark 1966). In some areas, creambush oceanspray is grown as a horticultural species for its dark green foliage, extremely long flowering period, and orange to dull red fall coloration (Sutton and Johnson 1974). It is one of the first shrubs to leaf out in the spring. The small, dry fruits were eaten by Native Americans. Figure 13—Rockspirea has smaller leaves and a Seedlings establish readily from seed or transplant more compact growth habit than creambush stock, but grow at only a moderate rate. Consequently, oceanspray; it often occurs on cliffs and rock they require protection from grass and forb competition outcrops.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 565 Chapter 22 Rosaceous Shrubs big sagebrush, pinyon, curlleaf mountain mahogany, Peraphyllum ramosissimum ______chaparral, aspen-lodgepole pine, spruce-fir, Douglas- fir, white fir, and ponderosa pine forests (Sutton and Indian apple Johnson 1974; USDA Forest Service 1937). Within its range it commonly occurs on dry to moderately dry Indian apple is a deciduous rounded shrub growing sandy or gravelly soils, rocky ridges, talus slopes, and to 6 ft (2 m) in height from numerous, gray-barked basal stems (fig. 14). Fascicles of simple, nearly sessile basalt outcrops (Alexander and others 1974; Hitchcock oblanceolate leaves 0.6 to 0.8 inch (15 to 20 mm) long, and others 1961; Welsh and others 1987). It is fre- and 0.2 to 0.4 inch (5 to 9 mm) wide are produced at the quently a codominant in climax communities, but has ends of short lateral spurs of the season. Leaves are been reported to occur as a pioneer species following glabrous above with a minute pubescence below. Per- disturbances in a white fir/Rocky Mountain maple/ fect, 5-merous, epigynous flowers appear with the rockspirea habitat type, where it persisted as the site leaves, singly or in clusters of two to five. Leaves may recovered (Alexander 1985). be dropped in midsummer, possibly in response to Cultural and management practices recommended water or temperature stress. Flowers are regular, for creambush oceanspray may be generally applied perfect, and fragrant with spreading pink to rose to rockspirea. King (1947) determined that although petals. The style and the 15 or more stamens inserted there are about 5,400,000 achenes per lb (12,000,000 on the edge of the hypanthium are exserted at anthe- per kg), only about 7 percent were sound. These resis. Fruit is a yellowish, bitter tasting, apple-like quired wet prechilling at 41 ∞F (5 ∞C) for 18 weeks to pome, 0.5 inch (12 mm) in diameter containing four release dormancy. Heat generated by wildfires may seeds. In Utah, Indian apple flowers from April to also stimulate germination. July. Fruits ripen and are gravity or animal dispersed The palatability and forage value of rockspirea gen- in July or August (Hitchcock and others 1961; Welsh erally receive low ratings (USDA Forest Service 1937). and others 1987). Kufeld and others (1973) reported its use by mule deer was moderate in fall and light during the remainder of Ecological Relationships and Distribution the year. It is used by birds and rabbits to some extent (Sutton and Johnson 1974; Van Dersal 1938). Based Peraphyllum is a monotypic genus of the subfamily on results of clipping studies in Oregon, Garrison Pomoideae distributed from southern and eastern (1953) recommended a maximum of 50 to 60 percent Oregon along the east side of the Sierra Nevadas use for sustained production. Present primarily on through central California and east to western Colorado summer ranges where other species receive preferen- and northwestern New Mexico at elevations ranging tial use, excessive grazing of rockspirea rarely occurs, from 3,000 to 9,000 ft (900 to 2,700 m) (Dayton 1931; and, at least on some western juniper/big sagebrush Ferguson 1983). It grows from the sagebrush/grass, habitat types of eastern Oregon, the species tends to pinyon-juniper, mountain brush, and oak/sagebrush increase when other species receive excessive use zones to the lower fringes of ponderosa pine forests (Ferguson 1983; Hopkins and Kovalchik 1983). Rockspirea thrives on dry slopes and unstable hillsides. It has high potential for use in revegetating disturbed sites (Johnston 1987; Stark 1966). Seeding practices and propagation of nursery stock are as described for oceanspray. Plants are occasionally planted as ornamentals because of their long flowering period, attractive foliage, and compact growth habit (Stark 1966; Sutton and Johnson 1974). Use of the species to date has been seriously limited by a lack of commercially available seed. Management of the species is also as described for creambush oceanspray. Little is known of the plant’s regenerative ability following use, fire, or other physical damage (McMurray 1987a). Sprouting capabilities have not been well documented. Some regeneration might occur from wind-dispersed seed or from soil seed banks.

Varieties and Ecotypes Figure 14—Indian apple is an intricately branched shrub with potential for use in low maintenance There are no releases. landscaping.

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(Hitchcock and others 1961; Welsh 1982). Chromosome protection for up to 3 years following planting. Indian number is n = 17 (McArthur and Sanderson 1985). apple may be entirely defoliated by grasshoppers when Historically, Indian apple appeared in the Conifer their populations are high. However, plants can re- Woodland Element of central Nevada’s Upper Mi- cover even after successive defoliations. ocene floras in an area where the Arcto-and Madro- Tertiary floras overlapped (Axelrod 1950). Varieties and Ecotypes

Plant Culture Indian apple accessions have received little testing. When selecting ecotypes for revegetation, emphasis Seed is harvested by hand picking or beating the should be placed on plant size, fruit production, growth fruits into containers. Fresh or dry fruits are macer- rate, and palatability. ated in a Dybvig or blender with water. Pulp that cannot be separated by flotation may be removed Physocarpus malvaceous ______from the seed by fanning after the pulp is dried. Approximately 6.5 to 10.3 lb (2.9 to 4.6 kg) of cleaned Mallowleaf Ninebark seed may be obtained from 100 lb (45 kg) of fruit (Swingle 1939). There are approximately 23,000 seeds Mallowleaf ninebark is an open, multiple-stemmed per lb (50,706 per kg) at 100 percent purity. Acceptable shrub 3 to 8 ft (1 to 2.5 m) tall (fig. 15) that spreads purity is 95 percent and germination is 50 percent for vegetatively from long rhizomes (Hitchcock and others seed purchases (see chapter 24). 1961; Welsh and other 1987). Roots are shallow and Seed requires a wet prechilling period of approxi- spreading. Bark is gray to red brown, shredding in mately 45 days at 38 ∞F (3 ∞C) to relieve dormancy thin layers. Leaves are alternate and currant- or (Belcher 1985; SEAM 1976; Smith 1974b). Germina- mallow-like with three to five palmate lobes. Leaves tion of seedlots tested by the Coeur d’Alene Forest are doubly crenate, nearly glabrous above, stellate Service Nursery ranged from 60 to 80 percent (SEAM pubescent to glabous below, and about 0.75 to 3.2 1976). Seed testing techniques have been described by inches (2.0 to 8.0 cm) long. Clusters of stellate white Belcher (1985). Seeds remain viable for up to 5 years flowers develop on the terminal corymbs. Flowers are if stored in a cool, dry, well ventilated container (Smith regular and perigynous. Fruits are paired, densely 1974). stellate, flattened follicles with ascending beaks that The existence of distinct ecotypes, possibly restricted dehisce along two sutures. Each follicle contains 1 to to specific soil types, makes selection of appropriate several tiny, hard, shiny, pyramiform seed each seed sources important. Seed should be fall planted on about 0.1 inch (2 mm) long (Gill and Pogge 1974a). medium textured, well-drained soils in adapted areas Mallowleaf ninebark flowers in May or June and sets (Stark 1966). Seeding may be accomplished by hand seed in August or September. Some fruits persist on planting or drilling with other shrubs, but separately the plant through the winter. from grass and forb species. Little seed is available commercially; production of bareroot or container planting stock may maximize the number of seedlings obtained (Ferguson 1983).

Uses and Management The forage value of Indian apple varies with accession and associated plant species. It is grazed by deer, particularly in fall. The fruits are used by birds and rodents. Indian apple can be incorporated into windbreak and conservation plantings on arid lands. The thick branching habit of the shrub offers cover for birds and other small animals in winter, even after the leaves have fallen. It also has potential for landscaping in arid areas, along roadways, and in recreation areas (Monsen and Davis 1985). It increases diversity on adapted sites. Indian apple is highly persistent once established, leading Blauer and others (1975) to suggest that it be used as a rootstock for related, but less tenacious species. Figure 15—Mallowleaf ninebark grows rapidly; Initial survival of Indian apple plantings is often its dense root systems stabilize soil on unstable quite low. Seedlings grow slowly and may require disturbances.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 567 Chapter 22 Rosaceous Shrubs

Like other shrubby members of the subfamily Plant Culture Spiraeoideae native to western North America, the chromosome number for mallowleaf ninebark is n = 9 Mallowleaf Ninebark seed are hand collected by (McArthur and Sanderson 1985). Species with n = 7 detaching the dried flower heads (Van Dersal 1938). and n = 9 occur among Asian taxa of the genus Seed is separated from debris using a barley debearder (McArthur and others 1983b). and an air screen fanning machine. There are approximately 756,000 seed per lb (1,666,700 per kg). Ecological Relationships and Distribution Acceptable purity for seed purchases is 98 percent and germination 40 percent (see chapter 24). Seed is Mallowleaf ninebark occurs east of the Cascades expensive and rarely available. Little is known of the from British Columbia and Alberta south through seed biology of this species. A high percentage of the Nevada, Utah, and Wyoming. It grows in association tiny seeds are normally not sound. Sound seeds are with mountain big sagebrush, aspen, and many coni- dormant and require a 77-day prechilling at 40 to 43 ∞F fer communities at elevations from 5,200 to 10,800 ft (4 to 6 ∞C) (Gill and Pogge 1974a; Marchant and (1,560 to 3,240 m) (Welsh 1982). It may be found on Sherlock 1984). dry open slopes, talus slopes, northern exposures, in Direct seeding is accomplished by broadcasting scattered to heavy timber, and on riparian sites (Davis seed onto a rough seedbed in fall and allowing natural 1952). Mallowleaf ninebark is adapted to deep, well- soil sloughing to cover it. Best results are obtained drained organic soils of medium texture and pH of when seed is planted separately from other species in 7.0 to 7.5 (Marchant and Sherlock 1984), but it will selected spots. Seedlings can establish with some grow on drier, sandy or rocky soils (Daubenmire and shade and limited herbaceous competition. Daubenmire 1968; Ferguson 1983). It is often abun- For bareroot nursery plantings, mallowleaf ninebark dant and highly productive. is fall seeded or artificially wet prechilled and spring Mallowleaf ninebark is frequently a member of seeded. Seed should be covered lightly and mulched. seral shrub communities that develop following log- Swingle (1939) estimated that a pound of seed would ging or wildfires in conifer forests (Olsen and Nagle yield about 30,000 usable seedlings. Seedlings are 1965; Owens 1982). The species is highly adapted to lifted as 1-0 to 2-0 stock depending on purchase speci- fire; it resprouts from the root crown or horizontal fications and growing conditions. Container stock may rhizomes. The more completely buried rhizomes have be propagated from germinants or cuttings. Hard- the greatest resprouting capacity (Bradley 1984). wood stem cuttings are easily propagated and fre- Owens (1982) and Merrill (1982) found that number of quently used. Container and bareroot stock develop plants and twig production increased on burned sites slowly, producing densely branched root systems. Both relative to unburned sites. In addition, recovery is container and bareroot seedlings establish readily enhanced by arrival of seed from undisturbed sites. (Monsen n.d.). The light seed is dispersed over long distances by wind and gravity (Young 1983). Uses and Management Seral shrub communities provide forage and cover for deer, elk, and other wildlife, but rapidly grow out of The value of mallowleaf ninebark as a forage plant reach of browsing animals. Overmature brushfields varies with season, location, class of game or livestock have been rejuvenated by prescribed burning, cutting, being considered, and availability of other vegetation. or topkilling tall shrubs with herbicides (Merrill 1982; It is generally considered to be more valuable on Mueggler 1966; Orme and Leege 1980). Control of ranges where better forage plants are depleted or brush species to promote conifer regeneration has also unavailable, and on drier sites (Noste and Bushey been accomplished with herbicides (Miller 1981; Miller 1987). Stands are rarely lost as a result of browsing. and Pope 1982a,b). The USDA Forest Service (1937) rated mallowleaf In addition to mallowleaf ninebark, several other ninebark as fair for sheep, fairly good for goats, and ninebark species occur within the Intermountain re- poor for cattle; Van Dersal (1938) rated it as good gion. Dwarf ninebark occurs in pinyon-juniper and browse for sheep, goats, and cattle. Kufeld and others related habitats of Idaho, Nevada, Colorado, and Utah. (1973) reported it receives light spring use and moder- Mountain ninebark is found on moist soils of aspen ate summer and fall use by mule deer. In cafeteria and coniferous forests from Nevada east to Wyoming style feeding trials with tame mule deer, Smith (1953) and south to Arizona and New Mexico. Pacific ninebark found mallowleaf ninebark was one of the most highly also grows on moist soils, but is restricted to California preferred native forages in the Logan, UT, area from and the area west of the Cascades in Oregon and May 1 to June 20. Because of its deciduous habit, it is Washington. A disjunct population occurs in northern of little value on winter ranges. New growth is more Idaho (Ferguson 1983; Hitchcock and others 1961). heavily browsed by deer for up to 3 years following burning (Noste and Bushey 1987).

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Mallowleaf ninebark has potential value as an erosion control plant. It grows rapidly, producing a network of rhizomes and a densely branched fibrous root system. It is a useful species for the restoration of high elevation acid minespoils in Nevada (Butterfield and Tueller 1980; Everett and others 1980). In spite of its rapid growth rate, mallowleaf ninebark receives little use in windbreak and farmstead plantings because of its bushy growth habit (Olsen and Nagle 1965). Mallowleaf ninebark has gained acceptance as an ornamental in some areas. It is a hardy, easily grown plant with attractive white flowers and dull to bright reddish brown leaf coloration (Gill and Pogge 1974a; Sutton and Johnson 1974). Plants that become too bushy can be pruned back. Once established, seedlings of mallowleaf ninebark grow rapidly compared to those of many other members of the Rosaceae. They compete successfully with other shrub species, but require some protection from grass competition for one or two growing seasons. Seedlings are not browsed heavily if alternative forage is available, but they may suffer predation by rodents and trampling by big game and livestock.

Potentilla fruticosa ______Shrubby Cinquefoil Shrubby cinquefoil, also known as bush cinquefoil, buckbrush, hardhack, ninebark, or yellow rose, is a Figure 16—Shrubby cinquefoil is a common ri- rather short lived, densely branched, round-topped to parian species in wet meadows at mid to upper low spreading shrub with spreading and fibrous elevations. roots. It produces white to bright yellow flowers throughout the summer, when adequate soil water is available (fig. 16) (USDA Forest Service 1937). The Hemisphere (Young and Young 1986). Shrubby cinque- shreddy brown bark of the main stems peels off in long foil is widely distributed in the Northern Hemisphere, strips. Leaves are pale to gray-green, leathery, 0.4 to ranging east from Alaska’s north slope to Greenland 1.2 inch (1 to 3 cm) long, and pinnately compound and Labrador and south to California, New Mexico, with five to seven narrow, often revolute, leaflets. Minnesota, and New Jersey. It also grows in Europe Leaves turn yellow in fall, but may be retained over and Asia (Hitchcock and Cronquist 1973; USDA For- winter. Flowers are 5-merous, complete, and solitary est Service 1937). In common with several mesic or in three to seven-flowered cymes. Each flower is shrubby North American members of the subfamily 0.75 to 1 inch (2 to 2.5 cm) in diameter with a saucer- Rosoideae, the diploid chromosome number of shrubby shaped hypanthium and five petals. There are 10 to 30 cinquefoil is 2n = 14 (McArthur and Sanderson 1985); stamens and numerous pistils. Fruits are golden tan, that of European plants is 2n = 28 (Elkington and teardrop-shaped achenes crowded on a hairy recep- Woodell 1963). Hitchcock and others (1961) reported tacle. Individual achenes are 0.04 to 0.08 inch (1 to 2 hybridization occurs within this genus, but hybridiza- mm) long with very thin seed coats covered with white tion of shrubby cinquefoil with other species has not hairs. Fruits mature and are dispersed continuously been observed (Elkington and Woodell 1963). by wind and gravity throughout the flowering season Shrubby cinquefoil is associated with cool climates (Hitchcock and others 1961). However, in Alberta, Canada, flowering apparently occurs in two flushes, and wet areas; it grows from plains and lower foot- one in May and the other in August (Scotter 1975). hills to above timberline over an elevation range of Fruits mature in mid to late summer and fall. 6,100 to 12,000 ft (1,850 to 3,600 m) (Belcher 1985). In the western United States shrubby cinquefoil is a Ecological Relationships and Distribution common summer range plant in wet subalpine meadows. It also grows near springs and marshes and along There are about 250 species of Potentilla occurring streambanks and floodplains in chaparral, sagebrush, in temperate to subarctic regions of the Northern aspen/lodgepole pine, ponderosa pine, Douglas-fir/white

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 569 Chapter 22 Rosaceous Shrubs fir, and spruce/fir forests (Sutton and Johnson 1974; Bareroot and container stock can be produced from Welsh 1982; Welsh and others 1987). It is most seed. Container stock started from germinants or abundant on areas with deep, wet, medium-textured small transplants requires a 3 to 5 month cropping soils and can withstand some salinity (Davis 1952; season (Landis and Simonich 1984). Wildings are an Stark 1966). alternative source of planting stock. The response of shrubby cinquefoil to burning de- Production of shrubby cinquefoil for the commercial pends on the season, site conditions, and fire intensity, ornamental market generally involves vegetative and may vary among populations. Plants may be propagation of horticultural varieties. Plants are ca- relatively unaffected to heavily impacted by burning pable of natural root sprouting following disturbances (Fischer and Clayton 1983; Mueggler and Stewart (Wasser 1982). Stem layering also occurs and may be 1980). Little damage is incurred by low intensity effective in spread and survival where inundation spring burns that fail to kill the root crown (Wasser occurs (Elkington and Woodell 1963). Vegetative propa- 1982), particularly on wet or rocky sites. Plants gation is accomplished by rooting softwood or regenerate rapidly by sprouting from root crowns, semihardwood stem cuttings in a greenhouse. Root rhizomes, and stem layering (Wasser 1982). Severe cuttings, layers, and crown divisions are also propa- burns occur when dense mature stands burn in late gated (Doran 1957; Mahlstede and Haber 1957; Toogood summer, particularly under droughty conditions. 1980). Some soil seed reserves may also be lost under such circumstances with the result that recovery occurs Uses and Management very slowly. Shrubby cinquefoil is an indicator of heavy grazing; Plant Culture it may become more abundant when other species are depleted. It is outcompeted by grasses and declines if Seed collection is difficult. Fruits are scattered over grazing is eliminated. Although its palatability is the crown of the plant and ripen unevenly during the often low due to its coarse, astringent foliage (Dayton growing season. Plants are not shade tolerant and 1931), game and livestock will use shrubby cinquefoil produce little seed when shaded (Scotter 1975). Little when other species are not available (Wasser 1982). seed is mature at any given time and hand collection Young plants are generally more palatable than ma- proceeds slowly. Consequently, seed is rarely avail- ture shrubs (Wasser 1982). In general, it is more able commercially (Ferguson 1983). Seed is cleaned palatable to sheep and goats than to cattle and horses with a debearder and fanning mill. It may be stored in (Sampson and Jesperson 1963). Because of its higher a warehouse for 5 years. Seventy percent is considered elevational distribution, it normally occurs on sum- an acceptable level for both purity and germination mer ranges, but where available in winter, it is some- for seed purchases (see chapter 24). Seed quality is times used because of its semi-evergreen habit and highly variable, but often rather low (Elkington and dominance in some areas (USDA Forest Service 1937). Woodell 1963; Wasser 1982). Kufeld and others (1973) found mule deer make Little information is available regarding germina- light use of shrubby cinquefoil in winter and take only tion requirements of shrubby cinquefoil seed. Vories trace amounts in spring and summer. Elk and white- (1981) reported the species is difficult to grow from tailed deer make some use of the plant during all seed. Belcher (1985) and SEAM (1976) found no germi- seasons. Bighorn sheep use it primarily in autumn nation pretreatment was required although a water (Van Dyke and others 1983). Seeds are consumed by soak at 86 ∞F (30 ∞C) for 18 hours did enhance germi- birds and small mammals (Wasser 1982). Palatability nation. McDonough (1969) applied an 11-minute soak of shrubby cinquefoil varies geographically and in sulfuric acid and obtained 73 to 87 percent seasonally. It is generally more palatable in spring germination at 90/72, 72/63, and 63/53 ∞F (32/22, 22/ and summer (Plummer and others 1968). Although it 17, and 17/12 ∞C). Light is required during incubation is avoided by big game in Alberta, it is unusually (Young and Young 1986). Tetrazolium and x-ray test- palatable in parts of Montana and Arizona. Livestock ing instructions were provided by Belcher (1985). also make greater use of it on scattered high altitude Direct seeding information is scarce, but seeding can summer ranges of the southwestern states, south- be successful on sites with good soil water. Babb (1959) western Montana, southeast Idaho, and Utah (Dayton recommended planting as soon as seed ripens. Plant- 1931; Ferguson 1983; USDA Forest Service 1937). ing before mid-July or in late fall was recommended by Plants are moderately tolerant of browsing, but they Wasser (1982). Seeding may be accomplished by drill- can be damaged by repeated grazing, particularly ing, broadcasting, or spot seeding on prepared sites during periods of drought when plants may become (Wasser 1982). Seeds are small and should be covered hedged and stunted and eventually lost (USDA Forest lightly, about 0.25 inches (6 mm) deep or less (Wasser Service 1937; Wasser 1982). 1982).

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Whether transplanted or grown from seed, shrubby Plant Culture cinquefoil is a useful soil stabilization species for disturbed sites with fairly wet soils. It is useful for American plum ranges from Manitoba through north- mine reclamation, roadway plantings, and game eastern Canada and south to New Mexico and north- range rehabilitation (Dittberner and Olsen 1983; western Florida (Grisez 1974; Hitchcock and Cronquist Fedkenheuer and others 1980; Ferguson and 1961). It occurs in the Rocky Mountain region and has Frischknecht 1985; Stark 1966; Wasser 1982). It is an been introduced into portions of the Intermountain excellent species for ornamental plantings because of area west of its native range. It will grow on deep soils its rapid growth rate, ease of cultivation, attractive in mountain brush, pinyon-juniper, and sagebrush semi-evergreen leaves, colorful flowers, and long grass habitat types receiving at least 16 inches (40 cm) flowering period. of annual precipitation (Plummer and others 1968; Although shrubby cinquefoil seedlings have been Thornburg 1982). Within its native range it is often a described as weak, they begin to grow rapidly once climax dominant or codominant, but it may be an early established and are better able to survive herbaceous seral species in some forested habitat types (Elkington competition than most other rose species. Plants are and Woodell 1963). It acts as a colonizer on unstable cold, but not drought tolerant (Sutton and Johnson sites along streams or rivers (Elkington and Woodell 1974). If planted in areas not heavily grazed by game 1963; Mueggler and Stewart 1980). Where introduced or livestock, they develop to maturity in about 5 years in the Intermountain region, it often grows in sandy to (Wasser 1982). loamy soils along ditch banks, drainage areas, and around home or farm buildings where supplemental Varieties and Ecotypes water is available. Plants are capable of withstanding some salinity. Considerable natural variability has been observed American plum does not establish well from direct in this species. Numerous ornamental varieties of seeding. However, it can be established from bareroot shrubby cinquefoil are marketed commercially. How- or container stock transplanted singly or in rows or ever, the species has received little testing for wild- clusters to meet planting objectives. Rootstocks are land uses. Populations with a rapid growth rate, utilized for propagation of domestic plum in northern spreading growth habit, and low palatability should climates (Grisez 1974). American plum grows well in be of greatest value for soil stabilization and mixtures with other woody species. Herbs with some streambank erosion control. Numerous showy flow- shade tolerance do well as understory species. ers, a long flowering period, low palatability, and American plum will sprout when burned, trampled, evergreen leaves are desirable characteristics for popu- or otherwise damaged. Spread also occurs from seed lations used for landscaping in recreation areas. transported by animals. The species has good drought and excellent cold tolerance. Prunus americana ______American Plum American or Pottawatami plum is an erect shrub or rarely a small tree. In the Great Plains it grows from 9 to 26 ft (3 to 8 m) in height (fig. 17) (Harrington 1964; Hitchcock and others 1961). In Utah plants are thicket-forming shrubs up to 16 ft (5 m) tall (Welsh and others 1987). Regeneration by rhizome sprouting is the main method of spread (McMurray 1987b; Welsh and others 1987). Branches are thornlike at the tip. Leaves are 1 to 2.8 inches (2.5 to 7 cm) long, elliptic to ovate or lanceolate and sharply long attenuate. They have pubescent, glandular petioles. Small umbels of white flowers are borne before the leaves on spur branchlets or axillary buds formed the previous season. Fruits are fleshy yellow to red drupes 0.8 inch (2 cm) in diameter, each with a single large stone. Some fruits may be hard and dry (Welsh and others Figure 17—In the Intermountain area, American 1987). Seed are dispersed by gravity and frugivorous plum is a thicket-forming shrub that spreads from mammals. rhizomes.

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Ecological Relationships and Distribution American plum flowers in April or May and fruits ripen in September or October (Grisez 1974; Plummer and others 1968). Fruits are harvested, conditioned, and stored as described for common chokecherry. Seed requires a 60 to 150 day wet prechilling to relieve dormancy (Grisez 1974). There are about 810 seed per lb (1,786 per kg) at 100 percent purity. Acceptable purity is 98 percent and germination 70 percent for seed purchases (see chapter 24).

Uses and Management American plum has been planted for many years in conservation plantings, windbreaks, and shelterbelts in the Midwest and Western States. It provides excellent escape, nesting, thermal, and travel cover for big game, livestock, and small mammals and birds. Leaves, twigs, and fruit are utilized by these animals. American plum can be an important component of erosion control plantings and low maintenance landscape projects. It has excellent potential for use in recreation areas, roadside rest areas, roadways, and administrative sites. It is also used as an aesthetic screen for wildlife in areas requiring thermal and concealment cover.

Varieties and Ecotypes Figure 18—Desert almond is a clonal shrub A number of American plum cultivars have been valuable for soil stabilization, cover, and as a selected, primarily for their fruiting characteristics. low water-use ornamental. There are no selections specifically developed for wildland use. Available material varies considerably in its range of adaptability and suckering habit. Stock adapted to the planting site and suitable for the separate clones within an area may vary by up to a planting goals should be acquired. month, ensuring that some flowers and fruits will survive late frosts (Kay and others 1977e). Fruits Prunus andersonii______ripen in June or July. The drupes resemble small, dry, fuzzy greenish-orange peaches and have a thin, rather Desert Almond or Anderson Peachbrush dry fleshy layer. The thick hard coat of the stone opens easily along one suture (Mozingo 1987). Fruits and Mozingo (1987) described desert almond as “one of seeds are dispersed by gravity and wildlife. the most beautiful and probably the most unappreci- ated shrubs in the western Great Basin.” Also known Ecological Relationships and Distribution as desert peach, Anderson almond, or Nevada wild almond; desert almond generally occurs as low grow- The native range of desert almond extends from ing, densely shrubby, clonal plants. Each clone may northeastern California southward along the eastern cover several acres. Individual stems are evidently slopes of the Sierra Nevada Mountains and eastward short lived, usually producing 6 to 8 annual rings across western and southern Nevada at elevations (Kay and others 1977f; Young and Evans 1973, 1974). from 3,450 to 7,400 ft (1,050 to 2,250 m) (Ferguson The plants have an average height of about 3 ft (1 m), 1983; Munz and Keck 1959; Young and Evans 1973). but range from 1.5 to 6.5 ft (0.5 to 2 m) in height It is a fairly drought tolerant species, most commonly (fig. 18). They are intricately branched and densely found scattered through big sagebrush and pinyon- thorny; each branch ends in a sharp spine. Narrow juniper communities on foothills, mesas, alluvial ter- leaves develop in fascicles on short, lateral branches. races, and in canyons. It is especially common on coarse Small pink to light red flowers appear in April to sandy or gravelly to rocky soil derived from decompos- June and cover the shrub. The flowering period for ing granite (Dayton 1931; Smith and others 1978).

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Plant Culture Prunus besseyi ______Seedling establishment apparently occurs only Bessey Cherry infrequently. Seedlings have been noted following removal of woody species in sagebrush/grass and Bessey cherry or western sand cherry is a low- pinyon-juniper sites (Young and Evans 1973, 1974); growing shrub with numerous spreading basal stems they develop from soil seedbanks or from off-site seed 1 to 5 ft (0.3 to 1.5 m) tall (fig. 19) (Vories 1981; Welsh transported by mammals (McMurray 1987c). Plants 1982). Leaf blades are 0.6 to 2 inches (1.5 to 5.0 cm) resprout vigorously following fire (Young and Evans long and 0.2 to 0.7 inch (5 to 18 mm) wide, elliptic to 1978b). Clones may survive into later seral stages of the oblanceolate, and acute to cuspidate apically. Leaves, community; maximum occurrence is on old burns that sepals, petals, hypanthia, and fruits are glabrous. have reached mid-successional stages (Koniak 1985). White flowers appear with the leaves in clusters of two Fruits are hand collected. Conditioning involves to four. The black fruits are 0.5 to 0.7 inch (12 to 18 macerating the fruits in a Dybvig or blender and mm) in diameter. Flowering occurs in April or May; floating the pulp. To relieve dormancy, Kay and others fruit ripening and dispersal occur from July to Sep- (1977f) recommended a 30 day wet prechilling; 60 days tember in Nebraska (Grisez 1974), and from August to were recommended by SEAM (1976). Seed should be September in Utah (Plummer and others 1968). fall planted to provide for field prechilling. Container and bareroot seedlings have been used successfully. Ecological Relationships and Distribution Planting stock can also be propagated from cuttings, although the success rate may not be high. Everett and Bessey cherry is native to the Great Plains and is others (1978a) recommended use of cuttings collected distributed from Manitoba south to Wyoming, during the period of seed maturation. Rooting re- Colorado, and Kansas (Grisez 1974). The species is quired approximately 6 weeks under a greenhouse commonly used as a dwarfing rootstock for other mist system. Prunus species. It has been introduced into the Intermountain region in wildlife and conservation Uses and Management plantings and by dispersal from orchards. Bessey cherry has been successfully established on sites Desert almond has good potential for use as a soil ranging from basin big sagebrush communities to stabilization species on disturbed sites due to its low, openings in ponderosa pine forests. Adkins (1980) dense growth form, clonal habit, and good establish- reported survival in conservation plantings in eastern ment. Everett (1980) obtained second year survival of Washington was 28 percent in the 9 to 12 inch (23 to 77 percent and 25 percent for containerized desert 30 cm) precipitation zone; survival ranged from 3 to almond seedlings planted on south- and north-facing 61 percent when plants were fully established. He roadcuts, respectively, near Reno, NV. Seedlings were concluded that in low precipitation areas Bessey tenacious, but tended to grow slowly. Smith and others cherry produced inadequate growth to provide good (1978) obtained 3-year survival of 73 percent when cover and was unable to compete successfully with desert peachbrush seedlings grown in containers cheatgrass and other annual and perennial weeds. 1.5 ft long (0.5 m) were transplanted in holes drilled into granitic bedrock near Mammoth Lake, California. They recommended the species for granitic sites within its range. Desert almond foliage is somewhat palatable, accessible, and probably seasonally important. Livestock and wildlife make some use of the species, particularly in spring or when summer storms induce new leaf production (Dayton 1931). The dense patches provide good cover for birds and other small animals. Desert almond has good potential as a low maintenance ornamental for roadsides, parks, and campgrounds because of its colorful flowers and long flowering period.

Varieties and Ecotypes No improved varieties are available for revegetation projects. Some variability among clones has been noted, but comparative testing of different popula- Figure 19—Native to the Great Plains, Bessey tions has been limited. Desert almond has been grown cherry has been used in wildlife and conservation as far north as Boise, ID, (Monsen n.d.). plantings in the Intermountain region.

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Plant Culture growth habit and several crooked stems commonly 3 to 13 (26) ft (1 to 4 [8] m) tall that grows in and east Seed collection, conditioning, storage, and planting of the Cascade Mountains (Hitchcock and others 1961). practices are generally as described for common The deep reddish-purple bark on young twigs of both chokecherry. Seed is often collected from stock main- varieties turns gray to dark brown on older branches tained at a nursery or from conservation plantings. and stems. The entire, alternate leaves are 1.2 to 2 Swingle (1939) found an average of 17 lb (7.6 kg) of inches (3 to 5 cm) long, glabrous to pubescent, and seed could be recovered from 100 lb (45 kg) of fruit oblong to obovate with crenulate to serrate margins. yielding 1,500 to 2,260 clean seed per lb (3,300 to Stipules are caducous and attenuate to linear. Corymbs 4,980 per kg). Babb (1959) obtained greatest germi- of 3 to 12 white insect-pollinated flowers (fig. 20) are produced from axillary buds on twigs of the previous nation following a 100 day wet prechilling at 41 ∞F season. Flowers are complete, regular, perigynous, and (5 ∞C) or 120 days at 33 ∞F (1 ∞C); he noted a high degree of variability in requirements among seed- 5-merous. Sepals are produced on a turbinate, decidu- lots. A 72 hour gibberellic acid soak (3.46 g Rootone F/ ous, disk-lined hypanthium that varies from glabrous to hairy. There are 12 to 20 exserted stamens and a 1000 g H2O) followed by wet prechilling for 20 days at 41 F (5 C) was recommended by SEAM (1976). single pistil. Only one of the two ovules develops. The ∞ ∞ fruit is a bitter tasting, red to black, one-seeded drupe Bessey cherry can be direct seeded. Seed should be from 0.3 to 0.5 inch (8 to 12 mm) in diameter. Bitter fall planted 0.5 to 1.0 inch (1.0 to 2.5 cm) deep on cherry flowers from April to June. Fruits ripen from seedbeds cleared of competition. The species can also July to September depending on elevation and local be established from container or bareroot stock (Monsen weather conditions (Grisez 1974; Plummer and others 1974). It can be propagated from buds, grafts, suckers, 1968). Fruits are dispersed in August or September. or root cuttings (Babb 1959). For windbreaks or con- Seed are often distributed by birds and mammals. servation plantings, a 4 ft (1.2 m) spacing is recommended (Cook 1981).

Uses and Management Monsen (1974) and Monsen and Christensen (1975) recommended use of Bessey cherry to provide good ground cover and soil stabilization on adapted sites in Utah and Idaho. It has been used extensively in the Midwest as a low growing erosion control and wildlife plant; it provides fruits and cover for birds and other small animals. Attractive flowers, fall leaf coloration, and its low, spreading growth habit make Bessey cherry a unique plant for landscaping. Rodents and birds often collect planted seed and emerging seedlings. Severe damage to planted Bessey cherry seedlings has resulted from rodent girdling (Brown and Martinsen 1959). Plants developing from seed or planting stock should be protected from excessive grazing and competition with grasses and forbs.

Varieties and Ecotypes Some selections have been developed for horticultural plantings, but no specific material has been developed for wildland uses.

Prunus emarginata ______Bitter Cherry Hitchcock and others (1961) recognized two varieties of bitter cherry: Prunus emarginata var. mollis, an arborescent variety growing to 42 ft (15 m) in height Figure 20—Bitter cherry flowers from April to June, and distributed west of the Cascade Mountains, and depending on elevation. Fruits are used by many var. emarginata, a more shrubby form with an open birds and mammals.

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Ecological Relationships and Distribution twigs and buds of plants growing on ridges and open slopes of winter ranges. The species is ranked as Bitter cherry occurs from the Pacific Coast of British highly valuable for elk in summer and fall (Kufeld Columbia east to northern Montana, and south to 1973). Mule deer use trace amounts in winter and southern California and southwestern New Mexico make moderate use of the plant during other seasons (Ferguson 1983; Hitchcock and others 1961). Its (Kufeld and others 1973). Its palatability has been elevational range varies with latitude. In the north it ranked as poor to fair for cattle and fair to fairly good grows from sea level to 3,000 ft (915 m); in the south- for sheep. Major livestock use occurs primarily in late west it occurs from 5,000 to 9,000 ft (1,525 to 2,745 m) summer or fall when other vegetation is no longer (USDA Forest Service 1937). available. In many areas dense thickets of bitter Bitter cherry is most frequently found in the upper cherry grow out of reach of grazing animals, but portion of the ponderosa pine belt. It is frequently provide good cover. Lower growing forms may be found on well drained, moderately fertile, rocky sites heavily browsed. Fruits are consumed by birds and such as ridge tops and southwest slopes in mountain mammals; rodents harvest the seed. brush communities. It is capable of resprouting Bitter cherry may be planted on shallow soils of following burning and often dominates other shrubs disturbed sites within its natural range to control in seral brushfields. Bitter cherry grows along water- erosion on roadways and other disturbances (Monsen courses in grasslands and sagebrush communities. and Davis 1985). Butterfield and Tueller (1980) and Everett and others (1980) concluded bitter cherry Plant Culture held considerable promise on acid mine spoils in Nevada. Establishment from seed or seedlings can be To maximize the number of viable seed obtained, hindered by trampling and grazing; the seedlings fruits should not be harvested until they are bright red develop slowly. Rodents consume seed and damage and fully mature (Grisez 1974). Fruits are collected by seedlings. Swihart and Yahner (1983) found rodents hand stripping or by placing tarps under trees to catch preferred Prunus species over the remaining 29 com- them following natural dispersal. They may also be mon windbreak and conservation species tested. For harvested with a mechanical tree shaker. Swingle cottontails, Prunus fell into a neutral to preferred (1939) reported that about 25 lb (11.4 kg) of seed can category. Seedlings develop slowly and must be pro- be obtained from 100 lb (45.5 kg) of fruit yielding tected from competition with more aggressive plants 1,800 to 5,471 seeds per lb (3,965 to 12,050 per kg). for up to 3 years following establishment. A Dybvig or commercial blender may be used to Bitter cherry spreads by root sprouts; new shoots macerate the fruit. All pulp and juice should be cleaned develop readily following burning. Leege and Hickey from the stones. Part of the pulp and some low quality (1971), working near the Lochsa River of northern seeds can be removed from the mixture by flotation, Idaho, found spring and fall burns increased bitter the rest by drying and fanning. Dried fruits are cleaned cherry sprouting about equally; mortality was slightly by hammermilling at a low speed, followed by fanning. higher in fall than in spring. Spring burning was Specific storage requirements have not been ascer- recommended for decreasing height of shrubs in seral tained for bitter cherry (Grisez 1974); seeds are com- brush fields and improving browse conditions for elk. monly stored as described for common chokecherry. Adequate regrowth occurred during the summer to Bitter cherry seed fill is generally low. Filled seeds provide fall forage. are difficult to germinate. Treatments to increase germination give inconsistent and generally poor results. Germination of treated seed is generally less Varieties and Ecotypes than 20 percent (Monsen and Davis 1985). Poor germi- There are no releases. nation has discouraged use of this species in direct seedings. The species can be intertransplanted using bareroot or container seedlings on favorable sites. Prunus fasciculata ______Seedlings grow remarkably well on dry and exposed sites with shallow soils where few other shrubs Desert Peachbrush establish. Wildings and root cuttings are alternative Desert peachbrush is a low growing, intricately methods of propagation. and divaricately branched deciduous shrub with a rounded growth form (fig. 21). Shoots may attain Uses and Management heights of 5 ft (1.5 m) and diameters of 8 ft (2.4 m). Plants have a deep taproot as well as a spreading Bitter cherry is frequently planted to provide browse fibrous root system. They are not rhizomatous for wildlife and livestock. Elk, deer, and moose eat (Harrington 1964; Welsh and others 1987).

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the stony endocarp is thin and dry and does need not be extracted. Leaves and sticks are separated by screening. Fruits are then dried and stored or seeded. There are about 4,500 seed per lb (9,900 per kg). Viability is generally good and is retained for 5 to 10 years of storage following harvest (Stevens and others 1981a). Seed retained in storage for 20 years exhibited fair viability. Acceptable germination for seed purchases is 90 percent and purity 70 percent (see chapter 24). Seed can be drilled or broadcast and covered. It must be placed 0.5 to 1 inch (1.25 to 2.5 cm) deep in a firm seedbed. Fall or winter seeding provides the wet prechilling period required to relieve dormancy. A strong, fast growing seedling is produced and good Figure 21—Desert peachbrush provides soil establishment is common if soil water conditions are stabilization and cover for wildlife. Because of its favorable. This species can also be propagated and unique branching habit, it has potential as a low outplanted quite easily as bareroot or container nurs- maintenance ornamental. ery stock (Monsen and Davis 1985). Considerable numbers of seedlings sometimes establish naturally from rodent caches.

Branches are grayish white, ending in pointed spines. Uses and Management The light green leaves are 0.1 to 0.2 inch (2.5 to 5 mm) Desert peachbrush is generally seeded in a mix with long and 0.04 to 0.2 inch (1 to 5 mm) wide, cuneate to other shrubs to improve diversity. California quail spatulate, and entire to few toothed at the apex. and Gambel quail, chukar, nongame birds, and small Flowers are small with five cream colored petals. Fruit mammals use it extensively for cover, nesting, and is a pubescent greenish-orange drupe with a rather forage. With the exception of pronghorn (Smith and dry fleshy layer. Beale 1980), livestock and big game make little use of the species. Although the shrubs are sometimes in- Ecological Relationships and Distribution fested with tent caterpillars, they persist in spite of defoliation. Desert peachbrush occurs in central and southern Desert peachbrush establishes and does well when Nevada, southwestern California, and western Arizona seeded on disturbed sites. Individual plants can cover on valley floors and foothills at elevations from 2,050 to large areas. Branches contact the soil surface over 5,800 ft (625 to 1,770 m) (Ferguson 1983; Welsh and most of the area and provide considerable ground others 1987). Desert peachbrush occurs in association cover and soil protection. Perennial forbs and grasses with chaparral, basin big sagebrush, Wyoming big emerge and develop beneath and adjacent to the crown. sagebrush, sand sage, blackbrush, creosotebush, Desert peachbrush can establish on roadcuts and fills, fourwing saltbush, quailbush, and lower pinyon-juni- in sand and gravel pits, and on mined areas. It also has per communities. It can be found on well drained to ornamental potential, especially for low maintenance heavy soils ranging from basic to neutral. Greatest landscaping projects. The intricate branching pat- area of occurrence is on sandy to rocky loam soils in tern, grayish-white stems, and light green leaves give depressions along drainage areas, intermittent the plant a unique appearance. streams, and creeks. Within these areas plants occur Seeded plants should be given 2 to 3 years to estab- as individuals or more typically in small groups, usu- lish and attain reasonable size prior to grazing. Once ally with a rather sparse understory. Following fire, established, this species is very hardy and can with- desert peachbrush sprouts readily and profusely from stand considerable use. the root crown. Seed dispersal and seedling estab- Desert peachbrush has performed well when planted lishment may be heavily dependent on rodents that on big sagebrush sites. It can be interseeded or gather and cache seed. intertransplanted in crested wheatgrass or native perennial herb seedings. Growth of established seed- Plant Culture lings is not suppressed by annual grasses. Because of its ability to persist, even with wildfires and invasions Harvestable seed crops are produced in about 1 of of annual weeds, increased study and use of desert every 5 years. Fruits are gathered by hand stripping or peachbrush to improve diversity of plantings on desert by beating them into a container. The pulp attached to shrublands within its range are warranted.

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Varieties and Ecotypes stems sprout from the rhizomes and the root crown. Leaves are alternate and elliptic, about 0.1 to 4 inches There are no subspecies or released varieties of (2.5 to 10 cm) long and finely serrate along the edges desert peachbrush. Little selection work has been (Welsh and others 1987). Purplish-red glands are attempted, but the species is being tested as a drought usually found near the tip of the petiole or along the tolerant rootstock for other Prunus species (Welsh and leaf base. The leaves are bright green above and paler others 1987). beneath. They turn bright red in early autumn. Elongate racemes of fragrant white flowers develop from April to June. Racemes are about 3 to 6 inches Prunus spinosa and Prunus (8 to 15 cm) long. The insect-pollinated flowers are tomentosa ______perfect; five petals spread from a gland-lined hypanthium. Only one of the two ovules normally devel- Blackthorn and Nanking Cherry ops. The fruits are red to purplish-black cherrylike drupes about 0.4 inch (1 cm) in diameter; each con- A number of introduced Prunus species from eastern tains a single seed with a stony endocarp (Grisez 1974; Europe and Asia are adapted to mountain brush, Welsh and others 1987). Seeds ripen from July to upper pinyon-juniper, and big sagebrush communi- September. Fruit production is greater during years ties of the Intermountain region. Blackthorn and of average or better rainfall and on plants growing in Nanking cherry, both Asian species, have been suc- full sunlight. cessfully planted in the Intermountain West. Black- Hitchcock and others (1961) recognized three vari- thorn grows to 13 ft (4 m) and Nanking cherry to 10 ft eties of chokecherry. Common chokecherry is a large (3 m) in height (Grisez 1974). Flowers of both species shrub or small tree growing to 50 ft (15 m) in height are generally numerous and very showy. Blackthorn with bright to deep red fruits. Black chokecherry is a produces a fleshy, dark purple, glabrous fruit; fruits small to medium sized shrub or small tree 13 (20) ft (4 of Nanking cherry are semifleshy, light purple, and [6] m) tall with dark, nearly black fruits and thick pubescent. leaves. Western chokecherry ranges from 7 to 13 Both species have been used in shelterbelt, wind- (20) ft (2 to 4 [6] m) tall with dark, nearly black fruits break, and conservation plantings. They have high and thick leaves. potential for use in and around recreational areas, administration sites, low maintenance landscaping Ecological Relationships and Distribution projects, and on disturbed sites such as roadcuts and fills or mine disturbances. Chokecherry occurs primarily in central and east- Both species transplant very well. However, they ern United States. It ranges from Saskatchewan to are fairly difficult to establish from direct seeding. Newfoundland and south to Kansas and North Caro- Fall planting is required as the seed require wet lina (Hitchcock and others 1961; USDA Forest Service prechilling. Good diversity can be provided by includ- 1937). Black chokecherry is distributed from eastern ing these species in mixtures with other shrubs. Grasses British Columbia to the Dakotas and southward and forbs grow well in association with both species, east of the Cascades from northern California to but seedlings should be protected from herbaceous New Mexico. Western chokecherry occurs primarily competition. They have excellent resistance to insects and grasshoppers, good cold tolerance, and are fairly drought tolerant. Livestock and big game make some use of twigs and leaves; fruits are taken by birds and rodents. Birds use these species for cover, nesting, and brood rearing. Seed and bareroot or container stock of both species may be obtained from commercial nurseries in the Midwest and Intermountain regions.

Prunus virginiana ______Chokecherry Chokecherry is a thicket-forming shrub or small tree with several deep roots and a well-developed, fibrous root system (fig. 22). Plants develop rhizomes Figure 22—Chokecherry provides palatable and that are 0.4 to 0.6 inch (1 to 1.5 cm) in diameter and nutritious browse that is used by many wildlife extend laterally about 3 ft (1 m) from the stems. New species.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 577 Chapter 22 Rosaceous Shrubs along the west coast from British Columbia south to can also be harmful. Belcher (1985) and Marchant and California (Hitchcock and others 1961; Welsh and Sherlock (1984) reported that air dried seed stored in others 1987). sealed containers at 37 to 41 ∞F (3 to 5 ∞C) will remain Chokecherry is distributed from low elevation foot- viable for 3 to 5 years. hills into mountainous areas at elevations from sea Chokecherry seeds generally have fairly high viabil- level to 9,000 ft (2,750 m) on sites with annual precipi- ity, but germination of untreated seed is generally tation of 12 to 30 inches (305 to 760 mm) (Thornburg quite low. About one-half of the viable seed will germi- 1982; USDA Forest Service 1937). It is often associ- nate in 69 days (Wasser 1982). Western chokecherry ated with sagebrush, mountain brush, pinyon-juni- generally germinates more readily than black per, and dry conifer types. It grows on moderately chokecherry (Monsen and Davis 1985). Wet prechilling acidic to basic or saline soils with silty to sandy can increase both the rate and percent of germination. textures that are moderately deep, fertile, and usually Seed is wet prechilled at 36 to 41 ∞F (20 to 23 ∞C) for 120 well drained. Scattered plants or small clumps may be to 180 days (Wasser 1982). Thoroughly soaking the found on talus slopes and other rather dry sites; larger seed prior to wet prechilling may improve effective- thickets develop along streams and around springs, ness of the treatment. Wet prechilling is adequate if ditches, or roadsides in semi-arid areas. In higher the stones have cracked, but the radicles have not yet elevation communities, common chokecherry tends to emerged. The rate of radicle emergence is normally occur on more open sites and along riparian areas. It rather low, but can be further reduced, if necessary, by is not adapted to high water tables or prolonged lowering the prechilling temperature once the stones flooding (Wasser 1982). have broken open (Grisez 1974; Wasser 1982). Frac- turing the endocarp, boiling, freezing, and various Plant Culture chemical treatments have provided inconsistent results and are sometimes detrimental (Grisez 1974). Chokecherry spreads by seed. It also spreads veg- The Association of Official Seed Analysts recommended etatively from buds produced on rhizomes and the root a 60 to 90 day wet prechill period at 37 to 41 F (3 to crown. However, it is not commonly propagated by ∞ 5 ∞C) for testing germination (Belcher 1985). They vegetative means. Leafy softwood cuttings can be rooted, also provided tetrazolium testing and radiographic but hardwood cuttings are not recommended (Hartmann procedures. and others 1990; Marchant and Sherlock 1984). Nonprechilled seed must be fall seeded, but artifi- The drupes fall soon after reaching maturity (Grisez cially prechilled seed may be planted in spring. Seed 1974). Birds and mammals harvest and spread the should be planted 0.5 to 1 inch (1.25 to 2.5 cm) deep in seed. Germinability may be increased after seeds a firm seedbed. Seed can be drill seeded or metered have passed through their digestive tracts (Grisez through a seed dribbler mounted on a crawler tractor. 1974). Fruits are collected either by hand picking or The former method has provided excellent results beating them into a canvas or into a container. Seed (Stevens 1992). This species can be seeded individu- harvested too early may have low germinability. If ally or as a component of a mixture of other shrubs and harvested too late, many seeds may already have nonaggressive forbs and grasses. Seedlings establish fallen and others will split while being cleaned, leav- most successfully when planted on sites that receive ing the embryo vulnerable to mechanical damage at least 15 inches (38 cm) of annual precipitation (Grisez 1974; Marchant and Sherlock 1984). Har- (Monsen and Davis 1985). They have often been planted vested fruits should not be exposed to heat as mold on drier sites with poor success. Seedlings of black may develop, and viability may be reduced. Fruits are and western chokecherry are not highly competitive, cleaned by macerating the fleshy material with a but become more so after 2 to 3 years (Monsen and Dybvig cleaner and flushing with water to remove the Davis 1985; Plummer and others 1968; Thornburg flesh from the seed. This is followed by drying and 1982). Once the root system becomes well established, further cleaning with a clipper cleaner. Fermentation plants can exist with other woody species or herba- may facilitate cleaning, but may adversely affect ceous vegetation and can withstand considerable germination (Grisez 1974). There are about 4,100 grazing. cleaned seed per lb (9,020 per kg) (see chapter 24). Bareroot stock of all chokecherry species may be For seed purchases, acceptable purity is 90 percent grown in the nursery. However, seed and new seed- and germination 70 percent. lings are often gathered by birds and small rodents. Seed may be sown immediately after cleaning One growing season usually provides stock of ad- without further drying. Short-term storage for a few equate size for transplanting (Shaw 1984). Although months requires only air drying. Optimal long-term initial growth of germinants is very rapid, the rate storage conditions have not been determined, but sometimes drops off considerably during the grow- warm, humid conditions result in rapid deterioration ing season; a 2 year cropping period is often necessary of seed quality. On the other hand, excessive drying

578 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

(Marchant and Sherlock 1984). Bareroot seedlings dried fruits to make pemmican cakes for winter use have a strong taproot system. Rhizome and new shoot (Welsh and others 1987). Tea was prepared from the development are initiated early in the season, often stems and bark (Craighead and others 1963). prior to lifting. New growth can easily be damaged All species of chokecherry are well adapted to fire, during lifting and planting, thus reducing the success particularly if burned when dormant or when soil of establishment (Monsen n.d.). Seed, germinants, or moisture is high (Anderson and Bailey 1980; Tisdale small transplants can be used to propagate container and Hironaka 1981; Young 1983). Although top growth stock. Seedling development requires a 3 to 5 month is easily killed, vigorous sprouting from rhizomes or cropping period (Landis and Simonich 1984). the root crown provides for rapid recovery following the initial decline (Leege and Hickey 1971; Noste and Uses and Management Bushey 1987). After the first postburn season, numbers of stems and coverage by the species can exceed Chokecherry has been extensively used to improve preburn levels for a number of years. Although estab- wildlife habitat. Plants typically form relatively open lishment from seed is quite rare on undisturbed sites, thickets that allow livestock and big game access to the species may be dependent on seedbanks for postfire abundant amounts of nutritious and palatable browse. regeneration, at least in riparian areas or higher Despite its deciduous nature, chokecherry maintains elevation communities (Kramer 1984; Wright and relatively high nutrient levels in late fall and winter others 1979; Young 1983). (Dietz 1972). Chokecherry provides fairly palatable browse for big game and is widely utilized by elk, Varieties and Ecotypes moose, and bighorn sheep (Plummer and others 1968); it is also the preferred deer browse on many winter There are no releases. ranges throughout the Intermountain west (Dietz 1972; Kufeld 1972; Kufeld and others 1973). Chokecherry receives its greatest use in spring and Purshia glandulosa ______fall, but elk and deer use it to some extent year around. Desert Bitterbrush It is moderately palatable to all classes of livestock, although it seems to be more heavily browsed by sheep Desert bitterbrush is an evergreen shrub resem- than cattle (USDA Forest Service 1937; Wasser 1982). bling Stansbury cliffrose and frequently mistaken for Fruits are taken by birds, bear, and many small it. Welch and others (1987) described desert bitter- mammals (Aune and Stivers 1985; Blauer and others brush as ranging from 5 to 9 ft (1.5 to 2.7 m) in height 1975; Gullion 1964; Noste and Bushey 1987; Wasser (fig. 23), generally upright, but sometimes low and 1982). Plants are productive and withstand moderate use. However, repeated heavy browsing results in highlining, hedging, and lower availability of forage. New sprouts may be killed by trampling or browsing. Grazing does not affect the next year’s fruit production; flowers are borne on twigs of the season. Chokecherry may be toxic if consumed in large amounts; leaves contain the glucoside amygdalin (Noste and Bushey 1987). Young shoots contain the greatest concentrations. Most problems occur in spring and early summer. Following fall frosts, the plants become essentially harmless. Livestock are not likely to consume fatal amounts of chokecherry except in areas of animal concentration. Chokecherry is extremely valuable for watershed and streambank stabilization, restoration of mine and roadway disturbances, landscape plantings in recreation areas, and increasing diversity in conservation and windbreak plantings (Ferguson 1983; Plummer and others 1968; Shaw and Cooper 1973; Wasser 1982). All chokecherries are edible, but have an extremely sharp and biting flavor if eaten before fully ripe. The Figure 23—Desert bitterbrush is a stable hybrid fruits make excellent wine, jellies, syrup, honey, fla- derivative of Stansbury cliffrose and antelope vorings, and perfume oil. Native Americans used the bitterbrush.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 579 Chapter 22 Rosaceous Shrubs spreading. Branchlets are prominently glandular. The Plant Culture glandular leaves are 0.1 to 0.4 inch (3 to 10 mm) long and up to 0.2 inch (5 mm) wide, cuneate, glabrous Seed production, harvesting, conditioning, and above, and slightly tomentose beneath. Petals are storage are similar for desert and antelope bitter- 0.2 to 0.3 inch (5 to 8 mm) long and creamy white to brush (Deitschman and others 1974c; Giunta and yellowish. Achenes are oblique and 0.8 inch (2 cm) others 1978). When ecotypes of both species are grown long, including the elongate style. Glandular ever- at common garden sites in Idaho and Utah, desert bitterbrush is normally smaller in stature than the green leaves and smaller, elongate achenes differen- erect antelope bitterbrush growth form. It produces tiate desert bitterbrush from antelope bitterbrush less seed than antelope bitterbrush and good seed (fig. 24). crops occur less frequently. There are approximately 21,000 seed per lb (46,200 per kg). Seed retain viability Ecological Relationships and Distribution for up to 25 years when stored under warehouse conditions (Stevens and Jorgensen 1994). For seed Desert bitterbrush is restricted to southern California, southern Nevada, southern Utah, and purchases recommended purity is 95 percent and northern Arizona (Koehler and Smith 1981; McArthur germination 90 percent (see chapter 24). A high percentage of desert bitterbrush achenes and others 1983b; Nord 1965; Stutz and Thomas 1964). Koehler and Smith (1981) and Stutz and fail to mature normally. Commercially harvestable Thomas (1964) considered it to be a hybrid derivative seed crops are produced once in every 2 to 6 years at native collection sites. Consequently, it is necessary of Stansbury cliffrose and antelope bitterbrush parentage that arose from hybridization and subsequent to harvest adequate quantities of achenes in good introgression between these two species (McArthur years to fill anticipated needs (Plummer and others 1968). Desert bitterbrush seed orchards have been and others 1983b). This hybridization is a relatively established at various locations in Utah and Idaho in recent development and populations remain geneti- an attempt to develop more consistent seed produc- cally distinct. Welch and others (1987) transferred tion centers. cliffrose, Cowania, to the genus Purshia because hy- Germination requirements of desert and antelope brids of cliffrose, antelope bitterbrush, and desert bitterbrush are similar. Both require short periods of bitterbrush occur where the distribution of these spe- wet prechilling to release dormancy. Desert bitter- cies overlap. brush seedlings are more difficult to establish than In southern Utah, desert bitterbrush grows those of antelope bitterbrush. They grow more slowly principally in the blackbrush type, and in pinyon- and attain maturity later when grown together in juniper extensions into this type, at elevations from common gardens. Seedlings also grow more slowly 3,460 to 4,470 ft (1,055 to 1,363 m) (Welsh 1987). It is under nursery conditions with regular irrigation. more common in southern Nevada and California Because of its slow growth, desert bitterbrush is than in Utah. It often occupies sites where cool and less competitive with annual weeds than antelope warm desert species interface and is often the domi- bitterbrush. nant or codominant shrub when present. Direct comparison of establishment attributes between the two species of bitterbrush may create a somewhat distorted picture. It is adapted to more xeric sites and when planted in such areas can perform well. It usually grows in areas receiving less than 14 inches (36 cm) of annual precipitation. Artificial plantings on these sites are often unsuccessful; the precipitation received may be too low or erratic to permit establishment. However, desert bitterbrush seedlings are relatively hardy and can survive adverse weather conditions if they once establish. Rodents quickly gather planted desert bitterbrush seed. Consequently, planting in late fall or winter is advised to reduce seed predation. Seeding directly into a weedy understory should be avoided. Seed- lings normally require 1 to 3 years to fully establish. Figure 24—Desert bitterbrush, (left) is Both bareroot and container grown seedlings estab- distinguished from antelope bitterbrush lish well and can be used to provide more rapid (right) by its glandular, evergreen leaves. establishment.

580 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

Uses and Management Purshia tridentata ______To date, most plantings of desert bitterbrush have Antelope Bitterbrush been made on upland pinyon-juniper sites where antelope bitterbrush is native (Davis 1983a). In gen- Antelope bitterbrush is an intricately branched, eral, antelope bitterbrush ecotypes respond better deciduous shrub varying in growth habit from low than desert bitterbrush at most of these planting decumbent spreading forms to upright arborescent sites. However, desert bitterbrush plants have estab- plants over 13 ft (4 m) in height (fig. 25) (Blauer and lished and persisted on antelope bitterbrush sites in others 1975; Welsh and others 1987). Leaves are central and northern Utah, southern Idaho, and south- alternate, simple, pinnatified or apically three-toothed central Idaho. Desert bitterbrush plantings have also and usually glandular. Flowers are borne on short been successfully established on pinyon-juniper/ spurs of previous year’s growth. They are numerous, blackbrush sites in southern Utah (Davis 1983a). white to yellow, and about 0.3 inch (8 mm) in diam- Desert bitterbrush exhibits important vegetative traits eter (Harrington 1964; Munz and Keck 1959; Welsh such as production of semi-evergreen leaves, ability to and others 1987). The fruit is a cartilaginous achene resprout following fire, and good drought tolerance. 0.2 to 0.5 inch (6 to 12 mm) long (Young and Evans Desert bitterbrush demonstrates adaptability to dis- 1983) with a persistent tapering style and a black turbances and can be used to seed infertile mine spoils pyriform seed (Blauer and others 1975). Plants are and roadways in adapted areas. Welch and others reported to be long lived. Nord (1965) found the aver- (1983a) reported that it reduces soil erosion and pro- age age of decumbent plants growing above 8,000 ft vides habitat for game and non-game animals. Roots (2,440 m) in northern California was 52 years; at may form symbiotic nitrogen-fixing relationships with the endophyte Frankia (Nelson 1983). Once established, desert bitterbrush grows well with herbaceous understory plants and can be managed to provide seasonal herbage, particularly winter forage for A big game and livestock. Welch and others (1983a) found desert bitterbrush plants in a southwestern Idaho common garden retained a significantly greater quantity of leaves in winter than collections of antelope bitterbrush and cliffrose. Crude protein content of the desert bitterbrush accessions exceeded that of the other two species. Palatability may be somewhat lower than that of antelope bitterbrush. However, where desert bitterbrush occurs, variety, quality, and quantity of other browse species are often low. As a result, considerable use is made of the species, particularly by deer and cattle. Its growth habit provides much needed shade and cover for small game and birds. Seeds are collected, eaten, and cached by rodents, birds, and reptiles. Insect populations associated with desert bitterbrush are often high, especially during flower- B ing and seed set.

Varieties and Ecotypes No selections of desert bitterbrush have been developed for release (Davis 1983a). Ecotypes vary in growth form from decumbent to upright. Populations or ecotypes having good drought tolerance (Davis 1983a), fire tolerance, winter leafiness (Welch and others 1983a), and nitrogen-fixation capabilities (Righetti and others 1983) have been identified. Because desert bitterbrush, antelope bitterbrush, and Stansbury cliffrose hybridize freely, it may be possible to com- Figure 25—(A) Mature antelope bitterbrush plant bine desired attributes of these three species. and (B) fruit.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 581 Chapter 22 Rosaceous Shrubs other locations the average age exceeded 150 years ponderosa pine forests. At lower elevations, in the (Ferguson 1983a; Nord 1959). southern portion of its range, it extends into blackbrush Populations of antelope bitterbrush are extremely communities. It may also be encountered on high variable. Giunta and others (1978) described ecotypes mountain exposures intermixed with aspen, limber with differing growth forms, fire tolerance, rates and pine, and Douglas-fir. It usually exists at elevations periods of growth, seed production, drought resis- ranging from 200 ft (60 m) in the Pacific Northwest to tance, nutritional qualities, heat tolerance, cold hardi- 11,500 ft (3,510 m) in the Sierra Nevada Mountains in ness, climatic adaptation, resistance to disease or areas receiving 12 to 25 inches (30 to 64 cm) of annual insect damage, palatability to game or livestock, ever- precipitation (Nord 1965; Stanton 1959; Tew 1983). green habit, vegetative reproduction, and shade toler- Antelope bitterbrush is often a component of mixed ance. Winward and Findley (1983) recognized similar shrub communities (Blaisdell 1953; Ferguson and variations in Oregon and California populations, but Medin 1983; Holmgren and Basile 1956), and is also noted marked differences in leaf size, shape, color, frequently a dominant in the northwestern part of its presence or absence of glands, flowering, and seed set. range (Franklin and Dyrness 1973; Mueggler and Many differences observed among populations are Stewart 1980; Nord 1965). Prostrate and semi-erect genetically influenced (Klemmedson and Ferguson growth forms of antelope bitterbrush occur most 1969; Nord 1959a; Plummer and others 1968; Wagle commonly as understory plants in forested communi- and Vlamis 1961). Ecotypes maintain their native ties or at high elevations (Nord 1965). Some low- growth habits and morphological characteristic when growing types are also encountered at low elevations planted over a wide range of sites. However, climatic and under dry conditions, but the upright, tree-like and edaphic conditions alter growth response to some forms usually dominate on more xeric sites (Nord extent (Davis 1983a; Edgerton and others 1983; Nord 1965). 1965; Shaw and Monsen 1983a). Some ecotypes are adapted to specific soil conditions. Prostrate growth forms occur on shallow, coarse Ecological Relationships and Distribution textured soils in central Idaho, while tall, erect forms grow on deep productive soils where basin big Cowania is derived from the neotropical Tertiary sagebrush may also occur (Tew 1983a). Plummer and flora that emerged in the Sierra Madre region of others (1968) reported that ecotypes from neutral or Mexico during the Tertiary epoch (Axelrod 1958). slightly acidic soils in Idaho are not adapted to calcare- McArthur and others (1983b) postulated that Purshia ous soils of central Utah. Davis (1983a) found that is an early derivative of Cowania that evolved in isola- only 24 of 259 accessions planted on pinyon-juniper tion. Chromosome number for both genera is n = 9 sites in Utah scored performance index values that (Alderfer 1977; McArthur and others 1983b; Sanderson would recommend their use in these areas. 1969). Present contacts between Purshia and Cowania Antelope bitterbrush ecotypes vary in their response are considered very recent; their partially separated to burning. Adams (1980) suggested that antelope flowering periods form a weak isolation barrier. Natu- bitterbrush evolved as a fire climax. Edgerton (1983) ral hybrids are found throughout most of Utah and reported rapid establishment of new seedlings fol- Nevada where Cowania and Purshia overlap (Stutz lowing fire in ponderosa pine/antelope bitterbrush and Thomas 1964). The hybrids backcross with the communities in the southern Cascade Mountains. parental stock, producing genetic introgression in Reestablishment from seed occurs erratically in re- both genera. Introgression is considered to be the sponse to climatic conditions and may be limited by major process contributing to genetic differentiation grazing, rodent activity, presence of annual weeds, of ecotypes (McArthur and others 1983b; Stebbins and other factors. Emergence from rodent caches is 1959; Stutz and Thomas 1964; Thomas 1957). Desert common in many areas. bitterbrush is a southern adaptive derivative result- Resprouting capabilities vary with ecotype. Some ing from this process. In addition, cliffrose charac- ecotypes in forested communities recover well by teristics are present in most populations of antelope resprouting after burning (Martin 1983; Rice 1983; bitterbrush that occur beyond the northern margins of Sherman and Chilcote 1972; Zlatnik 1999). Driver and cliffrose distribution (Stutz and Thomas 1964). others (1980) found 100 percent resprouting of plants Antelope bitterbrush occurs from the Cascade Moun- burned in a ponderosa pine/grass/antelope bitterbrush tains eastward through the Rocky Mountains and from habitat type. Antelope bitterbrush plants recovered British Columbia south to Arizona and New Mexico well from spring burning in ponderosa pine-antelope (Ferguson 1983; Hitchcock and others 1961; Koehler bitterbrush-pinegrass habitats, but little resprouting and Smith 1981; McArthur and others 1983b; Stutz followed fall burning when soil moisture was low and Thomas 1964). It grows in big sagebrush, pinyon- (Driver 1983). juniper, and mountain brush communities and

582 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

Billings (1952), working in the western Great Basin, in southern Idaho set and matured seed in 42 to 48 and Hormay (1943) in California found that antelope days in 1979 and 57 to 60 days in 1980. Blaisdell (1958) bitterbrush rarely resprouted following fire and sug- found the period of flowering and seed maturation was gested burning would permanently eradicate the spe- slightly longer (66 days) at sites near Dubois, ID. Seed cies. Blaisdell (1953) found that not all plants growing maturation dates can be predicted with reasonable on the Upper Snake River Plain in Idaho resprouted accuracy. Nord (1965) developed an equation using even after a light burn. Murray (1983) reexamined elevation and latitude to predict mean dates of seed sites studied by Blaisdell and reported no antelope maturation for stands in California. Actual harvesting bitterbrush recovery 43 years after burning. Bunting dates generally fell within 6 to 10 days of predicted (1985) and Bunting and others (1987) concluded that dates. the columnar ecotypes growing in central Idaho do Accurate determination of seed ripening dates is not recover from burning. At Wallsburg, UT, 80 to critical for commercial seed harvesters as seeds shat- 90 percent of the antelope bitterbrush plants burned ter quickly when mature. Achene development in a wildfire resprouted the first year following the progresses through a number of recognizable stages. fire. By the end of the third year mortality exceeded In southern Idaho developing achenes reach nearly 90 percent. This pattern is commonly observed follow- mature size by mid June (Shaw and Monsen 1983b). ing wildfires in antelope bitterbrush stands in Idaho During this period of rapid growth, the endosperm and Utah. remains milky. About 10 to 16 days prior to maturation, the endosperm changes from white to dark red Plant Culture (blood stage). Achenes dry rapidly during the last 3 to 5 days of this period. Antelope bitterbrush was perhaps the first native Achenes can wither and abort at any stage of devel- shrub used for wildlife habitat and rangeland res- opment. Late frosts, poor or inadequate soil water, and toration in the Intermountain region. Since the early insect damage are principal reasons for crop failure. 1940s antelope bitterbrush has been seeded in many Many failures occur during the “blood stage” just prior areas in the western United States. Holmgren and to achene maturation. Insects may destroy a large Basile (1956) reported that of approximately 50 spe- proportion of the seed crop (Basile and Ferguson 1964; cies tested in Idaho, antelope bitterbrush was the Basile and others 1964; Ferguson 1967; Ferguson and easiest to establish from direct seeding. others 1963; Furniss 1983; Furniss and Barr 1975). Nord (1965) reported that young plants growing Shaw and Monsen (1983b) reported 50 to 60 percent of on native sites usually begin producing seed in about all flowers that initially appeared failed to develop a 10 years. By contrast, plants grown in cultivated mature fruit in a southern Idaho seed orchard as a nurseries, or in native stands managed for seed pro- result of insect damage. duction, often begin producing seed when 2 to 4 years Prospective seed collection sites must be inspected of age. Latitude, elevation, season, and site conditions frequently, particularly when seeds are in the blood influence vegetative phenology and the date of seed stage to determine when and where harvestable crops maturation (Nord 1965). Leaf growth of antelope bitterbrush precedes the will be available. Harvesting is accomplished by strik- appearance of floral buds. At a common garden near ing branches with a paddle to dislodge seeds into Boise, ID, leaf growth of ecotypes from a wide geo- canvas or metal trays (Giunta and others 1978; Nord graphic range was initiated between March 17 and 23 1967). Most native sites produce 150 to 200 lb of seed (Shaw and Monsen 1983b). Leaves reached mature per acre (168 to 224 kg per ha) although production size by about mid April when flower buds were begin- may approach 500 lb per acre (560 kg per ha) (Nord ning to develop. Leader growth of the different 1965). ecotypes was initiated during the first week of May, Few seed nurseries have been established on agri- coinciding with anthesis. Most stem elongation occurred cultural land. Cultivated plantings should be estab- in June and early July. Stem growth rate diminished lished on or near native sites. Plantings should be in July. Little elongation occurred in late summer or located in areas protected from high wind; flowers fall. All five ecotypes examined responded similarly. are cross pollinated by insects and seed shatter Climatic conditions favorable to good seed produc- easily (Blauer and others 1975). In southern Idaho 6 to tion antedate the crop by at least 1 year; flower buds 10 year old plants of an upright ecotype planted at 6 ft develop primarily on second year wood (Nord 1965). (1.8 m) spacings produced more seed per acre than Plants that receive favorable moisture produce good plants planted at 6 by 12 or 12 by 12 ft (1.8 by 3.7 or leader growth and increased root reserves that sup- 3.7 by 3.7 m) spacings. However, as plants attained port seed production the following year (McCarty and mature stature, those at 6 ft (1.8 m) spacings became Price 1942) (fig. 25b). so overgrown that seed harvesting became impracti- Shaw and Monsen (1983b) reported that accessions cal. Depending on growth habit, shrubs should be of antelope bitterbrush growing in a common garden

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 583 Chapter 22 Rosaceous Shrubs spaced 12 to 15 ft (3.7 to 4.6 m) apart to permit good seed through seeding mechanisms. Seeds require a 2 seed production and facilitate site maintenance and week wet prechilling to release dormancy, thus they seed harvesting. should be fall or winter seeded. Maintaining a stand of native herbs as an under- Recommended seeding rates for antelope bitter- story is a practical approach for controlling weeds and brush vary considerably. The shrub may be drill seeded, supporting predatory insects. Field cultivated plants interseeded in rows at various interspacings, or hand appear to benefit from cultural treatments, but re- planted in clearings or selected spots. Hubbard (1964) sponse to fertilizer, water, and pesticides is not known. recommended drilling at a rate of 3 lb per acre (1.4 kg Native stands of antelope bitterbrush can also be per ha). Extensive pinyon-juniper chainings have been managed for seed production. Plants respond favor- interseeded using a Hansen seed dribbler. Rows are ably to pruning, thinning, weed control, and protec- usually spaced 8 to 12 ft (2.4 to 3.7 m) apart when tion from grazing. mechanical interseeders are used (Stevens and others Annual collections of antelope bitterbrush from west- 1981b). Between 5 and 20 seeds are usually dispensed ern rangelands have sometimes exceeded 40,000 lb per linear ft (16 to 66 per m) in each interseeded row. (18,182 kg). Considerable seed is harvested annually At a rate of 20 seeds per ft (66 per m), approximately from most Intermountain and Northwestern States. 87,000 seeds are planted per acre (214,974 per ha), if Demand varies considerably from year to year. Areas rows are spaced 10 ft (3 m) apart. This equates to 3 to in south-central Idaho and northern California tend 5 lb per acre (3.4 to 5.6 kg per ha) depending on the seed to produce more consistent seed crops than most sites lot planted. within the Intermountain region. More seed is usually Hand seeding has been used extensively to plant harvested from areas in central Idaho and the Pacific steep or inaccessible slopes. Herbaceous vegetation Northwest than is used in those regions. However, must be cleared to enhance shrub seedling survival. seed collected from central Utah and eastern Nevada Clearings amid annual or perennial vegetation is generally not sufficient to meet regional demands. should be between 2.5 to 3.3 ft (0.75 to 1 m) square Likewise, the quantity of seed harvested from arid to effectively reduce competition. Hand seeding ante- regions and decumbent growth forms growing in lope bitterbrush and other shrub seed into depres- ponderosa pine and aspen communities is generally sions created by uprooting pinyon-juniper trees fol- inadequate. lowing anchor chaining is also a successful planting Seed are easily hand harvested. Effective mechani- technique. cal harvesters have not been developed (Nord and Rodents actively gather planted seed (Evans and others 1967). Air-dried collections are cleaned by others 1983; Young and Evans 1978a,b), and forage on first screening to remove sticks and debris. The small seedlings. Rodent damage can be extremely husks are then detached with a barley debearder, serious and limit planting success. This problem can Dybvig, or other scarifier. Remaining material is again be reduced by planting late in fall after rodents hiber- screened to separate seed from the detached husks nate. Coating seeds with a combination of Arasan and (Giunta and others 1978). At 100 percent purity there Endrin eliminates nearly all rodent predation (Medin are approximately 15,000 seed per lb (33,000 per kg). and Ferguson 1980), but the toxic effects of Endrin on Seed are reasonably hard and are not easily broken, other animals have eliminated its use. Everett and fractured, or damaged when cleaned or planted. Rec- Stevens (1981) found thiourea treatments reduced ommended purity for seed purchases is 95 percent and deer mouse predation. However, the compound is germination 90 percent (see chapter 24). now classified as a carcinogen. No other chemical has Cleaned seed can be stored under warehouse condi- been developed that effectively diminishes rodent tions for extended periods without loss of viability. predation. Afterripening improves germination. Stevens and oth- Evans and others (1983) and Reichman and Oberstein ers (1981a) reported a significant increase (79 to 86 (1977) reported that size and depth of seed caches percent) in germination between the second and third influenced detection by rodents. Rodents locate seed year of storage. Seed germination remained unchanged primarily by olfactory cues (Howard and Cole 1967; after 20 years of warehouse storage, but decreased to Johnson and Jorgensen 1981), thus edaphic condi- 24 percent after 25 years. Antelope bitterbrush seed tions can affect seed discovery (Johnson and Jorgensen can usually be stored without suffering extensive 1981). Rodents were better able to locate antelope insect damage, but storage areas must be kept rodent bitterbrush seeds when two or more seeds were planted free. together. Rodents failed to locate individual seeds Antelope bitterbrush seed are large, smooth, and placed 0.4 inch (1 cm) deep, but removed 75 percent of fairly heavy. Consequently, they are easily carried the seeds if two seeds were placed together at this through most seeders. When planted in mixtures, ante- depth. Predation increased to 98 percent when more lope bitterbrush can be used to carry other light-weight than 10 seeds were planted together. Evans and others

584 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

(1983) found that planting depth affected seed detec- and young plants are often browsed heavily. Project tion. Rodents consumed all seeds placed on the soil seedings or plantings should be large enough to dissi- surface or planted at a depth of 0.4 inch (1 cm), but pate animal browsing or establishment may be ad- only about 50 percent of seeds planted between 0.8 versely affected. and 1.6 inch (2 and 4 cm) deep. Available cover also strongly influences seed preda- Uses and Management tion. Evans and others (1983) reported significantly less rodent predation of seed caches in burned areas The broad genetic base of antelope bitterbrush and lacking plant cover than in unburned sites with con- its wide geographical distribution (Stutz and Thomas siderable vegetative cover. Similar responses have 1964; Thomas 1957), provide considerable opportu- been noted in seedings on mine and roadway distur- nity to promote the use of natural and artificial popu- bances. Rodents apparently cache antelope bitter- lations and hybrids with specific traits. Few other brush seeds in small openings, but are not likely to shrub species are so diverse. venture onto exposed sites for extended foraging. Antelope bitterbrush is widely regarded as an im- Planting success of range and wildlife seedings in portant browse plant for big game and domestic live- different plant communities also indicates consider- stock. It has been planted extensively in big game able differences in rodent foraging habits (Basile and habitat improvement projects throughout the Inter- Holmgren 1957; Casebeer 1954; Holmgren and Basile mountain and Pacific Northwest regions to provide a 1959; Nord 1965). Success has generally been better forage base in areas where game and livestock use in pinyon-juniper communities cleared of trees by may be seasonally heavy. Numerous seeding and anchor chaining than on burned antelope bitterbrush/ transplanting projects have been conducted to im- bunchgrass ranges infested with cheatgrass. prove sites where shrub populations have been de- Booth (1980) and Evans and others (1983) found pleted or eliminated. Antelope bitterbrush grows pretreating antelope bitterbrush seeds with a fungi- well with a wide number of native understory species cide reduced damping-off problems and ultimately and other shrubs and is often planted in mixtures to increased the number of successfully established seed- reestablish plant diversity. It forms important asso- lings. Cutworms and wireworms can also destroy ciations with various conifers, mountain brush spe- newly developing seedlings of antelope bitterbrush cies, and bunchgrasses. However, in certain areas (Hubbard 1956), but outbreaks are usually limited to this plant exists with few other species and may be isolated situations. the dominant plant encountered. In either situation, Compared to most shrub and many herb species, reestablishing the shrub improves wildlife habitat, antelope bitterbrush seedlings establish quite suc- watershed resources, and community and structural cessfully even under harsh circumstances. Conse- diversity. quently, the shrub can be used to revegetate road- A wide array of ecotypes exists within the broad ways, mine disturbances, and other infertile sites. If geographic range of antelope bitterbrush. Numerous protected from browsing, young plants grow rapidly ecotypes have been propagated and planted at various and provide considerable forage and ground cover study sites in the western United States. Some collec- within 3 to 5 years. tions exhibit adaptation to a rather broad range of site Antelope bitterbrush is often interseeded with a conditions; others require rather specific edaphic and combination of grasses and broadleaf herbs. However, climatic conditions. An upright growth form from Monsen and Shaw (1983c) reported that intermediate Sanpete County, UT, for example, performed much wheatgrass and crested wheatgrass planted as an better than any other source when seeded on a broad understory can significantly reduce or minimize natu- array of Utah sites (Davis 1983). A natural hybrid ral seedling recruitment of this shrub. It is essential of antelope bitterbrush and Stansbury cliffrose dis- that the shrub be seeded with less competitive herbs. played better survival, persistence, and herbage yield Interseeding antelope bitterbrush onto sites domi- than most other antelope bitterbrush ecotypes when nated by cheatgrass is not recommended unless reme- seeded on a number of Utah sites. dial treatments are employed to reduce or replace the Planting failures sometimes result when antelope cheatgrass understory. bitterbrush ecotypes are planted on similar parent Bareroot or container seedlings are frequently used materials but at elevations about 2,000 ft (610 m) to circumvent seed predation by rodents and speed higher than the native stands. In addition, ecotypes establishment. Dormant or carefully hardened seed- have failed to persist if sources from strongly basic soils lings should be planted before native antelope bitter- are planted on acidic sites, or vice versa. Selective brush shrubs at or near the planting site break grazing by livestock, big game, rodents, and insects has dormancy (Carpenter 1983). Establishment success is also eliminated specific ecotypes when they were planted usually quite high. Antelope bitterbrush seedlings in small test plots (Edgerton and others 1983; Giunta

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 585 Chapter 22 Rosaceous Shrubs and others 1978). If nonfire-tolerant ecotypes are planted Varieties and Ecotypes on sites subjected to burning, entire stands can be eliminated in a single fire. Land managers should be Numerous populations of antelope bitterbrush with very selective when considering the use of seed collec- distinctive attributes have been recognized and are tions obtained from areas differing from the prepared commercially harvested and sold (table 2). Ecotypes planting site in edaphic or climatic conditions. from central Utah are widely used for big game habitat Antelope bitterbrush is particularly well adapted to improvement, but are best adapted to calcareous soils. harsh sites and frequently acts as a pioneer species. Decumbent growth forms commonly found as an under- Consequently, it is frequently seeded on mines, road- story in lodgepole pine or ponderosa pine forests are ways, and similar sites. Decumbent and semi-erect used in revegetation of forested sites. Upright ecotypes growth forms can be used to control erosion; they from southern Oregon and central Utah are used to provide a dense, spreading ground cover capable of stabilize sandy soils and restrict human activity. spreading by stem layering. ‘Lassen’ antelope bitterbrush, the only release to Plants may form symbiotic nitrogen-fixing relation- date, is a robust, upright ecotype that produces and ships with the endophyte Frankia (Nelson 1983). maintains moderate quantities of overwintering Antelope bitterbrush is more frequently nodulated leaves that provide a source of protein for fall and than other rosaceous nitrogen-fixing species, but the winter grazing (Shaw and Monsen 1986; Welch and degree of nodulation varies considerably among others 1983a). This selection is native to Lassen County, antelope bitterbrush populations. The significance of CA, where it grows on coarse textured, granitic soils. nitrogen fixation in arid land rosaceous species is It has done well throughout the Pacific Northwest and poorly understood. the Intermountain regions when planted on sandy, Fire-tolerant ecotypes have been utilized to plant slightly acidic or neutral soils in areas receiving 14 to areas subjected to frequent burning. Most ecotypes 18 inches (36 to 46 cm) of annual rainfall. Seedlings planted for this purpose retain their resprouting char- are vigorous and grow rapidly. acteristics when planted on adapted sites. Conse- quently, selection programs have been conducted to Rosa woodsii ultramontana ______identify populations that are capable of resprouting. Antelope bitterbrush possesses many attributes that Woods Rose contribute to its widespread use. Seed are large, easy to collect, process, store, and plant with conventional Hitchcock and others (1961) and Welsh and others equipment. Sufficient seed crops are produced with (1987) described Woods rose as a deciduous shrub with frequent regularity to provide satisfactory amounts of a growth habit ranging from nearly prostrate to up- seed for most project plantings. In addition, nursery right and height from 3.3 to 10 ft (1.0 to 3.0 m). Its root and greenhouse-grown transplants are easily pro- system is dense and fibrous. Thick briar patches of duced and establish well when outplanted. Commer- arching branches are formed by heavy suckering from cial growers produce good quality stock, and with spreading rhizome systems. Branches are armed with careful planning, land managers can normally con- straight to curved infrastipular spines. They may tract production of adapted ecotypes. produce internodal prickles, but are occasionally un- The broad array of available growth forms provides armed. The alternate leaves are odd pinnate with five a useful assemblage of material for planting distur- to nine ovate to obovate or elliptic leaflets. The leaves bances. With proper selection, adapted ecotypes can are 0.6 to 5.1 inches (1.5 to 13 cm) long and 0.2 to 1 inch be used to provide windbreaks, serve as conservation (0.4 to 2.5 cm) wide, singly to doubly serrate, and and wildlife plantings, or be used for more formal pubescent to glandular or rarely stipitate glandular. maintenance landscape purposes. Different growth The fragrant flowers are solitary or in few-flowered corymbs or corymbiform cymes that terminate lateral forms can be used to enhance recreational sites, sum- branches of the season (fig. 26). Each flower has five mer homes, and parkways. persistent erect to spreading sepals and five white to Because antelope bitterbrush is a major wildlife deep rose petals. Numerous stamens are inserted on a species and establishes quite well, some grasslands disk lining the throat of the calyx. Carpels are numer- and shrublands have been converted to antelope bit- ous. The “hips” are globose to ellipsoidal and 0.2 to terbrush by artificial seedings. Some plantings have 0.5 inch (6 to 12 mm) in diameter. They turn a bright failed when the shrub was not well suited to the orange red at maturity. Each contains 15 to 30 straw problem area. Attempts to convert wildland sites to colored to white achenes 0.1 to 0.2 inch (3 to 5 mm) long antelope bitterbrush stands should be carefully with long stiff hairs along one side. evaluated. Unless adapted ecotypes of antelope bitter- Woods rose blooms from late spring to summer brush exist, more suitable species should be selected depending on water availability. Fruits ripen in late for planting. fall and remain on the shrub through winter (Blauer and others 1975; Hitchcock and others 1961).

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Table 2—Principal ecotypes of antelope bitterbrush used in wildland restoration projects.

Mean annual Growth Areas of Availability Ecotype/origin Elevation precipitation form Attributes adaptability Seed a Transplantsb References c feet inches Boise Front 2,800 17 Erect Highly productive, Neutral to acidic 4 3 2,3,4 Ada Co., ID good seedling soils, south-central vigor, excellent Idaho cover

Bryce Canyon 7,000 16 Spreading to Leafy, highly Colorado River 2 1 3,4 Garfield Co., UT semi-erect available forage, drainage, but resilient to heavy widely adapted browsing, excellent ground cover

Columbia Basin, 12-16 Spreading Excellent drought Columbia River, 2,3 eastern Washington tolerance, highly western and available forage, central Idaho good ground cover

Elko 5,200 14 Semi-erect Excellent drought Northern Nevada, 3 2 3,4 Elko Co., NV tolerance, persistent western Utah, responds well follow- big sagebrush ing fires, widely and pinyon-juniper adapted sites

‘Lassen’ 4,200 14 Erect Good winter Mid-elevations, 4 2 5 Lassen Co., CA leaf retention, northern California seedling vigor, eastern Oregon, palatability southern Idaho

Maybell 6,000 14 Spreading to Stem layering, Specifically 3 2 3 Moffat Co., CO semi-erect adapted to sandy adapted to soils, good sandy soils ground cover

Panguitch Lake 8,250 20 Decumbent Cold tolerant, Spruce-fir,aspen, 2 1 Garfield Co., UT excellent summer and mountain brush forage, resists types, exposed sites, browsing, excel- windblown exposures lent ground cover

Pinto 6,000 14 Upright Drought tolerant, Adapted 2 1 Washington Co., UT productive, good throughout the forage, excellent pinyon-juniper wildlife habitat, type within the consistently good Great Basin seed crop

Sanpete 5,000 14 Erect, upright Excellent seedling Most widely adapted 4 2 1,3,4 Sanpete Co., UT vigor, highly ecotype tested through- productive, good out the pinyon-juniper seed producer, type within the Inter- persistent mountain area

Shoshone 4,200 10 Decumbent Drought tolerant, Southern Idaho, 1 1 Lincoln Co., ID spreader, adapted basalt soils to rocky lava substrates

Snowville 5,100 15 Semi-erect Cover and forage Adapted to big 3 2 Box Elder Co., UT for big game and sagebrush and upland birds,excel- pinyon-juniper lent seed producer communities

Starvation 8,124 18 Decumbent, Highly palatable, Adapted to 2 2 3,4 Canyon spreading excellent summer high elevations, Wasatch Co., UT forage, resistant aspen openings to heavy grazing

Salt Lake 5,525 17 Erect, upright Good seed Well-drained 3 2 3,4 Salt Lake Co., UT producer, soils, mountain forage available brush and upper elevations of pinyon-juniper communities

aSeed availability: (1) Erratic producer - limited amounts available; (2) Fair to good producer - limited amounts collected; (3) Fair to good producer - moderate amounts collected most years; (4) Fair to good producer - large collection sites, seed usually available; (5) Good producer - seed normally available. bTransplant availability: (1) Produced only on request; (2) Produced mainly on request; (3) Produced mainly on request, some speculative rearing; (4) Produced for speculative sales. cReferences: (1) Davis (1983a); (2) Edgerton and others (1983); (3) McArthur and others (1983b); (4) Plummer and others (1968); (5) Shaw and Monsen (1986).

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variation in height, spininess, vegetative spread, and drought tolerance. This variety occurs from plains to alpine areas along marshes, lake shores, and in other riparian communities of the sagebrush, juniper, mountain brush, aspen, and spruce-fir zones. Plants may be found in well drained, wet to dry, loamy to sandy soils on all aspects, and in moderate shade to full sunlight. Woods rose grows at elevations from 2,800 to 11,000 ft (850 to 3,355 m) (Welsh and others 1987), generally in areas receiving 16 to 20 inches (41 to 51 cm) of annual precipitation (Monsen and Davis 1985; Wasser 1982). Woods rose invades disturbed areas along roadways, eroded drainages, fence rows, irrigation canals, and abandoned fields. It is intolerant of poor drainage, high water tables, and prolonged flooding (Wasser 1982). Ecotypes vary in drought tolerance; flowering is more adversely affected by drought than is rhizome growth. Woods rose resprouts following fire. However, intense fires or repeated burns can damage or kill the shallow root crown. Annual growth may increase following light to moderate burns (Wasser 1982).

Figure 26—Woods rose flowers in late spring or Plant Culture early summer. Fruits ripen in fall and contain about 15 to 30 achenes. Woods rose begins flowering when plants are 2 to 4 years old. Plants growing in full sunlight produce more flowers than those growing in shade. Good seed crops are produced at about 2 year intervals (Gill and Pogge 1974b). The hips of Woods rose are harvested by Ecological Relationships and Distribution hand as soon as they ripen and before the flesh begins to soften. Achenes are extracted by macerating the Rosa is a difficult genus taxonomically. It occurs hips in water in a blender or Dybvig and floating off throughout the Northern Hemisphere, and exhibits a the pulp and empty achenes. Achenes should be dried high degree of intraspecific variability. There are thoroughly prior to storage. Acceptable viability is re- approximately 115 species of Rosa in temperate North tained for 2 to 4 years by sealing the dry achenes in America (Young and Young 1986). About 10 of these containers at 34 to 38 ∞F (1 to 3 ∞C) (Gill and Pogge occur in the Rocky Mountains and most will hybridize 1974b). There are about 45,300 seed per lb (99,868 per (Hitchcock and others 1961). Chromosome number for kg) at 100 percent purity. Acceptable germination for this group is n = 7. The genus Rosa was represented seed purchases is 95 percent and purity 70 percent in the western American element of the Arcto-Ter- (see chapter 24). tiary Flora (Axelrod 1950). Dormancy of Woods rose seed is caused by inhibitors The native range of Woods rose extends from British in the seed coverings and mechanical restriction of the Columbia to southern California and east to pericarp wall (Jackson and Blundell 1963). In nature, Saskatchewan, Minnesota, Missouri, and Texas. There consumption and digestion of seeds by animals dis- are two varieties. The eastern variety, Rosa woodsii perse the seed and alleviates dormancy (Morris and var. woodsii occurs on plains and prairies as far west others 1962; Shaw 1974). Overall seed viability is as eastern Montana. Plants are about 3.3 ft (1 m) tall decreased by digestion, but dormancy of unharmed with crowded leaves. The leaflets are 0.4 to 0.8 inch seed is reduced. A wet prechilling at 34 to 41 ∞F (1 to (1 to 2 cm) long and gland tipped. The cordilleran 4 ∞C) for 30 to 365 days is recommended to relieve variety, Rosa woodsii var. ultramontana, covers the dormancy of planted seeds (Gill and Pogge 1974b). western portion of the species range. It is taller than Warm pretreatment for 60 days at room temperature var. woodsii, often reaching heights of 3.3 to 6.6 preceding the wet prechilling may also be helpful. (9.9) ft (1 to 2 [3] m). Its leaves are also larger, up to Hot water and acid treatments have been used prior to 2 inches (5 cm) long by 1 inch (2.5 cm) wide with coarse wet prechilling with mixed results (Gill and Pogge teeth that are not gland tipped. It exhibits tremendous 1974b).

588 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

In the field, seed may be broadcast or drilled at a rate Livestock, game animals, and rodent populations may of about 0.5 to 1.0 lb per acre (0.6 to 1.1 kg per ha). It require control during the period of establishment. should be covered with 0.25 to 0.75 inch (0.6 to 1.9 cm) Rodents consume planted seed and young seedlings of soil. Seed can be mixed with other shrub seed, but sometimes suffer serious debarking and girdling it should be seeded separately from grasses; seedlings (Spencer 1958; Wasser 1982). Plants are sometimes are only poorly to moderately competitive with herbs. attacked by leaf spots, rusts, downy mildews, blights, Seed may also be spot planted in areas cleared of and cankers. Periodic close use, mowing, or burning competition. Due to variability in seed dormancy among under safe conditions may improve the appearance seed collections, and the extremely long wet prechilling and stimulate production of plants along roadways or requirements, results of seedings have been quite in recreation areas. erratic. Woods rose is an extremely useful plant on dis- Transplanting, although more expensive than di- turbed sites such as roadcuts and fills, gullies, ripar- rect seeding, is often recommended, at least for small ian areas, and mining disturbances where soil sta- planting sites. Bareroot stock is usually lifted after bilization is essential (Monsen 1974, 1975; Plummer one growing season. Seedlings produce an extensive and others 1968). At the Leviathan Mine in western root system during the first growing season in the Nevada, Woods rose and mountain snowberry pro- nursery. They may initiate new rhizome growth prior vided the highest third year survival and cover of the to lifting. Rhizome growth is initiated early in spring. species tested (Everett and others 1980). Suckers were New rhizomes are easily damaged during lifting and produced the third year. Plants survived in spite of a outplanting. Large bareroot seedlings with shoots 2 to low soil pH (4.1) and low nitrogen. Liming and other 3 ft (0.6 to 0.9 m) tall are grown at some nurseries. amendments were recommended. Although natural These are useful on unstable or dry sites; the large root seedlings may invade disturbances, they are easily system provides anchoring and a greater water ab- buried by soil sloughing. However, seedlings do de- sorption area, and the long shoots are not easily buried velop quite rapidly if established from rooted cuttings by soil sloughing. Augers or other special planting or nursery stock. The fibrous, rapidly developing root tools may be required to properly plant these seed- system and rhizomes provide soil stabilization. Rhi- lings. Wildings may be dug when dormant and kept in zomatous spread has been noted by the third year cold storage for later transplanting. They should be after transplanting, augmenting the stabilization pro- handled and planted like bare root stock. cess. In addition, the brushy habit, thorniness, and Containerized seedlings may be grown as tublings low palatability of some ecotypes permit development or as larger plants. Seedlings may be started from of impenetrable thickets that reduce livestock and seed, small transplants, or cuttings (Landis and wildlife trampling on unstable sites. Many ecotypes, Simonich 1984). They require at least a 3 to 5 month however, are grazed rather heavily. Planting them in cropping time. a mixture with other shrubs is advised to take advan- Hardwood cuttings collected when dormant may be tage of differing growth habits and microsite require- stored at 34 ∞F (4 ∞C) and rooted in the nursery in areas ments and maximize the diversity obtained. with mild climates or under a greenhouse mist system Woods rose obtained the highest assessment of suit- in other areas. Softwood cuttings of partially mature ability of 18 shrubs tested on forest roads in north- wood root quickly (10 to 14 days) under a mist system. western Montana (Hungerford 1984). The assessment Rooted cuttings may be transplanted in fall, over- was based on growth, spread, vigor, survival, flower- wintered in a shadehouse or cold frame, or transferred ing, and regeneration. Overall survival after 4 years to a nursery bed (Gill and Pogge 1974b). Everett and was 62 percent with similar survival on south, west, others (1978a) recommended the use of hardwood and east aspects. Plants spread by root sprouting and rather than softwood cuttings. They used a mist sys- were indistinguishable after 4 years. The plant was tem and 0.8 percent indole-3-butyric acid to induce recommended for use in this area to hold soil against rooting in a greenhouse. Layers or suckers may also be wind and water erosion in open to light shade and in propagated in the nursery or greenhouse and field dry to moist soil. Everett (1980) found Woods rose and planted when adequate root systems have developed antelope bitterbrush were among the best species (Hartmann and Kester 1990). tested for revegetating roadcuts in Nevada. After 3 years, Woods rose survival remained high and the Uses and Management species was among the best for both north and south slopes when ranked by the amount of cover produced. Because of its wide geographic distribution and The species also exhibited the highest survival of revegetation values, Woods rose is a candidate for use species tested on forest roadcuts at 14 locations on two in a wide variety of planting situations. High survival forests in eastern Washington (Tiedemann and others rates of transplanted seedlings make Woods rose a 1976). Overall survival of Woods rose was 56 percent desirable and reliable addition to many projects.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 589 Chapter 22 Rosaceous Shrubs on the Colville National Forest and 63 percent on the and Utah at moderate elevations. Rosa acicularis, a Okanogan National Forest. Plants were generally of circumpolar species, extends south from Idaho and rather low vigor and on these sites may have required Montana to northern New Mexico. Rosa gymnocarpa fertilization. Successful planting required stems tall occurs from sea level to 6,000 ft (1,830 m) from British enough that they were not buried by raveling. Columbia to northwestern Montana, south to the Woods rose is used by big game during most seasons; Sierra Nevada Mountains of California and the Blue it generally receives heaviest use in early spring and and Ochoco Mountains of eastern Oregon (Hitchcock fall (Blauer and others 1975; Wasser 1982). Plants are and others 1961). It is a larger shrub with a more erect moderately grazing tolerant, particularly on moist and less thicket-like growth habit than Woods rose. soils. Palatability of leaves and stems varies with Uses and culture of these species are generally as ecotype, density of prickles, and availability of other described for Woods rose. forage. Wood rose is rated as valuable in winter and highly valuable in summer and fall for elk (Kufeld 1973). Mule deer make light use of it in winter and fall Rubus leucodermis ______(Kufeld and others 1973). In a cafeteria style study, Black Raspberry, Blackcap Smith (1953) found Woods rose was one of the most preferred shrubs for mule deer throughout the sum- Black raspberry or blackcap is a sprawling, decidu- mer (May 1 to September 30) in northern Utah. ous shrub with arching biennial canes ranging from Woods rose provides food and excellent nesting, 3 to 10 ft (1 to 3 m) in length (Hitchcock and others escape, and thermal cover for birds and small mam- 1961). Prickles of the leaves, petioles, and stems are mals during all seasons (Blauer and others 1975; retrorsely curved. The pinnate leaves are petiolate Thornburg 1982; Young and Young 1986). Although with linear, caducous stipules, and three to five not highly preferred, the hips remain on the shrubs doubly serrate leaflets. Leaflets are green and nearly well into the winter and provide an excellent energy glabrous above and white tomentose beneath. They source for many species when the ground is covered are about 2.4 to 5.6 inches (6 to 14 cm) long; the with snow (Mace and Bissell 1986; Monsen and Davis terminal leaflet is 1 to 3 inches (3 to 7.5 cm) long. 1985; Plummer and others 1968; Welch and Andrus Perfect, 5-merous white flowers are produced in 1977). racemes in groups of two to seven. Stamens and pistils Nutritive value of Woods rose varies considerably by are numerous. Fruits consist of aggregates of black season. In the Black Hills of South Dakota, protein drupelets on a convex receptacle. Each drupelet con- value varied from 4.9 percent in summer to 12.0 tains a hard seed. Flower and fruit development occur percent in spring for stems and from 5.7 percent in fall to 16.4 percent in spring for leaves (Dietz 1972). Rose unevenly through the summer. Seed is dispersed by hips have high digestibility and are moderately high birds and mammals. in crude protein in winter; they provide food for many birds and rodents. Ecological Relationships and Distribution Rose hips have been used as preserves and as an Rubus is distributed worldwide, but is most common ingredient in herbal tea and candy. They are high in vitamin C. American Indians used the bark, roots, and in temperate regions of the Northern Hemisphere. stems to produce a dressing for wounds, sores, and There are several hundred species. Many taxa are burns. Roots were used for treating diarrhea (Craighead interfertile; most are polymorphic, and a few are and others 1963). Woods rose has high ornamental apomictic. Rubus belongs to the subfamily Rosoideae value for recreation areas, parks, and campgrounds. and has chromosome number n = 7 (McArthur and The dense growth habit can also be used to direct Sanderson 1985). pedestrian traffic and provide low maintenance Black raspberry is distributed from British Columbia shrubbery. The attractive fruits, flowers, and fall east to Montana and south to California and New coloration also contribute to its landscape value. Mexico (Davis 1952). Although commonly a plant of moist environments such as riparian zones and forest Varieties and Ecotypes openings, black raspberry also exists on dry, open slopes of mountain brush communities and on talus No populations of Woods rose have been released for slopes or roadbanks. It spreads by layering and rhi- commercial production. Tremendous variability occurs zome sprouting as well as from seed and may be one in growth habit and many other characteristics. Some of only a few species growing on some rocky sites. accessions have a wide range of adaptability. Several additional Rosa species are used in wildlife Uses and Management plantings. Rosa nutkana is distributed from Alaska south to northern California and the Blue Mountain Ferguson (1983) considered black raspberry the most region of Oregon. It also extends south into Colorado likely of the Rubus species to be used for revegetation

590 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs purposes. It is easily propagated vegetatively and transplants readily. On roadcuts, drainageways, or other disturbed sites it grows rapidly, spreads vegetatively, and provides soil stabilization as well as dense cover and food for birds and other small animals. Black raspberry grows rapidly and is used only lightly as a summer forage by big game and livestock, thus its establishment is normally not hindered by excessive use.

Plant Culture Seed dormancy and cultural practices for black raspberry are similar to those described for thimbleberry. Time requirements for wet prechilling and scarification have not been determined. Fruits are collected when they are black and detach easily from the receptacle. Use of planting stock rather than seed on disturbed sites may provide better and more rapid success; these sites offer poor conditions for seed germination and early seedling establishment. Plant- ing stock may be grown from seed. Root cuttings, suckers, layers, stem cuttings, and leaf-bud cuttings are easily started (Hartmann and Kester 1990). Root cuttings may be treated as wildings, particularly on moist sites. Seedlings grow fairly rapidly, but are sensitive to dense competition and shading.

Varieties and Ecotypes

There are no releases. Figure 27—Thimbleberry spreads by seed or resprouting in natural forest openings and disturbed Rubus parviflorus ______areas. Thimbleberry Ecological Relationships and Distribution Thimbleberry is a low growing, rhizomatous, thicket- forming shrub. It is unarmed, deciduous, and 2 to 7 ft Thimbleberry is distributed from Alaska east to (0.5 to 2.0 m) tall (Hitchcock and others 1961). Bark of Ontario and south to Chihuahua, Mexico. It grows in the main stems is gray to brown and shreddy. Stems, all western States from elevations near sea level on petioles, and leaves are unarmed and stipitate glandu- the Pacific Coast to near 10,000 ft (3,050 m) in Colorado lar. The large leaves are long petioled with membra- (Harrington 1964; Sutton and Johnson 1974; USDA nous lanceolate stipules. They are alternate, simple, Forest Service 1937). Thimbleberry forms dense palmately lobed, and veined from a cordate base. patches in natural forest openings or disturbed areas, Leaves are 2 to 6 inches (5 to 15 cm) long and 0.6 to 2 but it is also fairly common in areas shaded by forest inches (1.5 to 5 cm) wide. The showy, white flowers are canopies, brushfields, or woody riparian plants, par- perfect and 5-merous, in groups of two to seven in flat- ticularly in the southern portion of its range. It is a topped cymes or terminal corymbs (fig. 27). Stamens particularly common understory species in the humid and pistils are numerous. Fruits are approximately forests of the Pacific Northwest. 0.5 to 1 inch (1.2 to 2.5 cm) wide aggregates of coherent Thimbleberry grows on moist, sandy loam soils rich red drupelets that are thinly fleshy and almost dry at in organic matter, but may also be found on rocky talus maturity. Fruit aggregates form over a conical recep- slopes, road banks, and creek bottoms. It regenerates tacle, thus the common name “thimbleberry.” Each rapidly from rhizome and root crown sprouting follow- drupelet contains a single hard beige to pink seed ing burning or logging (Cholewa and Johnson 1983; about 0.1 inch (2 mm) long. Flowering and fruit ripen- Leege and Hickey 1971; Mueggler 1966; Noste and ing continue through the summer. Seeds are dispersed Bushey 1987). Buried thimbleberry seeds have been by birds and mammals. reported in a number of forest seed bank studies

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(Kramer 1984). Although Kramer (1984) found low thimbleberry in fall and moderate amounts in sum- densities and low viability of thimbleberry seeds in mer. Thimbleberry is ranked as highly valuable and soil banks for three forest habitat types in central valuable for elk in summer and fall, respectively Idaho, he concluded that the rhizomatous nature of (Kufeld 1973). Palatability for livestock varies with the species would allow even isolated seedlings to geographic location and plant association. In gen- spread over significant areas. eral, it is considered worthless to poor for cattle and fair for sheep, although it may be heavily used if other Plant Culture vegetation is scarce or unpalatable (USDA Forest Service 1937). Good seed crops are produced almost every year but Primary values of thimbleberrry in revegetation fruits are difficult to collect in large quantities as they plantings are to provide soil stabilization and thickets are scattered over the crown of the plant and ripen for wildlife food and cover on disturbed forest sites, unevenly. Ripe fruits are bright red and detach easily sand dune areas, and streambanks. Seedlings are from the receptacle. They are dispersed by gravity and not likely to be browsed heavily unless other more by animals that consume them. palatable vegetation is unavailable. They are suscep- Fruits are hand collected. Seeds are extracted by tible to white fly and spider mite infestations (Marchant and Sherlock 1984). macerating the fruits in water in a Dybvig or blender Thimbleberry is a common plant of seral shrub and floating off the pulp. Remaining debris and light communities; it resprouts following logging or burn- seed are removed by drying and fanning. Dried seeds ing, and establishes in natural forest gaps. Burned may be stored at 41 ∞F (5 ∞C) for several years (Brinkman plants resprout from root crowns and rhizomes. In- 1974h). tense fires might destroy these structures, but permit Dormant embryos and an impermeable endocarp germination of seed banked in the soil (Noste and prevent rapid seed germination. In nature, dormancy Bushey 1987). Gratkowski (1978), Leege and Hickey is overcome by warm pretreatments and wet prechilling (1971), and Miller and Kidd (1983), discussed the occurring between early summer seed fall and spring response of thimbleberry to burning and herbicide germination, although seeds are capable of remaining treatments, practices used to kill shrubs and permit viable in the soil for more than 1 year if germination conifer regeneration or rejuvenate overgrown shrub conditions are not met (Brinkman 1974h). A 2-hour fields to improve browse availability for wildlife. soak in sulphuric acid or a 7-day soak in 1 percent sodium hypochlorite are commonly used germination Varieties and Ecotypes pretreatments. However, results of acid scarification have been erratic (Marchant and Sherlock 1984). None. Swingle (1939) and Babb (1959) recommended planting the seed immediately after it is harvested. Sum- Sorbus scopulina ______mer seeding in the nursery provides warm pretreatments and wet prechilling that relieve both blocks to Greene’s Mountain Ash germination for many Rubus species (Babb 1959; Brinkman 1974h). A 90-day warm pretreatment at 68 Greene’s mountain ash is an attractive semi-erect shrub, branching from the base and forming thickets to 86 ∞F (20 to 30 ∞C), followed by a 90 day wet by suckering or layering (fig. 28). Bark of younger prechilling at 41 ∞F (5 ∞C) in the laboratory, prepares seed for spring sowing. stems is smooth and yellowish gray becoming grayish Thimbleberry is easily established from seed-grown red on older stems. Young growth is sparingly to bareroot or container stock. Plants are also easily densely grayish strigillose pilose. Winter buds are glutinous and white hairy to glabrous. The alternate propagated from layers or suckers. Sprigging, the col- deciduous leaves are glossy dark green above and pale lection and planting of dormant rhizomes, provides a and glabrous beneath, turning a showy red orange in means of quickly obtaining large quantities of plants. autumn. They are pinnately compound with 7 to 13 serrate leaflets each, short cuneate to acute at the Uses and Management base, and acuminate at the tip. Stipules are green, Rubus was ranked first among 18 genera of shrubs membranous, and caducous to persistent. The inflo- in terms of the number of wildlife species using them rescence is a 70 to 200 flowered flat-topped corymb. The creamy-white flowers are perfect, 5-merous, and (Martin and others 1951; Robinette 1972). The drupelets are sought by birds and many other animals; regular (Preston 1968). Hypanthium lobes are trian- leaves are used to varying degrees in summer. The gular, persistent, and white hairy externally. There are 15 to 20 stamens and two to five carpels and dense sprawling plants provide cover for small animals during all seasons. Kufeld and others (1973) styles. Fruits are fleshy scarlet to orange pomes that reported that mule deer take trace amounts of dry to a purplish color. Each contains four seeds

592 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs

to 38 ∞F (1 to 3 ∞C) (Harris and Stein 1974). Properly stored seeds remain viable for 2 to 8 years. Seed requires wet prechilling for germination. A 72- hour soak in gibberellic acid (3.46 g Rootone-F/1000cc/ distilled H2O) followed by a 90-day wet prechilling at 41 ∞F (51 ∞C) was recommended by SEAM (1976) and Belcher (1985). Seed can also be given a 115-day warm pretreatment at 77 ∞F (25 ∞C) followed by a 75-day wet prechilling at 41 ∞F (5 ∞C), although exact requirements vary widely between and within seedlots. Labo- ratory germination tests require extended incubation periods. Excised embryo tests and tetrazolium chloride tests are frequently used alternatives; both can be completed rapidly (see Belcher 1985). Results of both Figure 28—Greene’s mountain ash fruits persist tests are generally higher than laboratory germina- into winter and are consumed by birds and other tion test results (Harris and Stein 1974). wildlife species. Greene’s mountain ash may be established by sowing untreated seed in late summer to provide warm and cold wet pretreatment periods. Laboratory prechilled seed may be spring planted. Seed should be (Harris and Stein 1974; Hitchcock and others 1961; covered lightly. Even when wet prechilled, seed lots Welsh and others 1987). The shrubs flower from May often germinate in the field over a 2 to 3 year period. to July and fruits mature from August through Octo- Seed should be planted in areas where it will be ber. Many fruits remain on the plant through winter. protected from competing vegetation. Seeds are spread primarily by birds. Production of planting stock may be necessary if seed supplies are limited or more rapid establishment Ecological Relationships and Distribution is desired. Container and bareroot stock are usually Approximately 50 species of Sorbus are distributed grown from seed. Bareroot stock develops slowly and through temperate and subarctic areas of the North- may require two growing seasons before lifting. Propa- ern Hemisphere (Hitchcock and others 1961). The gation by cuttings and layering is difficult. Wildings chromosome number n = 17 is typical of woody may be used as alternative sources of planting mate- Pomoideae genera in the western United States rial if only small quantities of plants are needed. (McArthur and Sanderson 1985). Greene’s mountain Cook (1981) recommended use of Greene’s mountain ash is distributed from Alaska east to Alberta and ash on adapted sites below 6,500 ft (2,000 m) in south to California, New Mexico, and the Dakotas Wyoming, and suggested that it be planted at 4 ft (Welsh 1982). It grows in openings in conifer forests (1.2 m) spacings. Young plants develop rather slowly from foothills to subalpine or alpine sites at 4,000 to and require protection from competition with grasses 9,000 ft (1,200 to 2,700 m) (Davis 1952; Stark 1966; and forbs. Seedlings are tenacious and evidently not Welsh 1982). Greene’s mountain ash grows in sun or very vulnerable to insect and disease problems. They partial shade on well-drained, neutral to acidic soils are taken by deer (USDA Forest Service 1948). forming dense patches or intermixed with other shrub species (Van Dersal 1938). It is common in mountain Uses and Management brush communities of forest openings, riparian areas, and on disturbed sites. The persistent fruits of Greene’s mountain ash are taken by bears and birds in the fall and into the winter. Plant Culture Moose often browse the shrubs heavily in winter. Twigs are eaten by game and livestock. Dayton (1931) Fruit should be gathered immediately when ripe to reported that Greene’s mountain ash produces good prevent losses to birds. Seeds are collected by hand or sheep feed. It provides good cover and soil stabilization shaken onto a canvas. Maceration of fruits in water in on steep sites within its range as it spreads to form a Dybvig or commercial blender separates seeds from the remainder of the fruit. The pulp contains germina- dense thickets. As an ornamental, it offers attractive tion inhibitors and is removed by flotation and by foliage, flowers, and fruits for mass plantings in full fanning and screening the dried seed and trash sunlight or partial shade. The foliage and berries are (Hartmann and Kester 1990; Heit 1968). Seed quality brightly colored in fall, and clusters of berries remain is often low. Seed should be dried to a 6 to 8 percent on the plant through the winter (Sutton and Johnson water content and stored dry in sealed containers at 34 1974).

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 593 Chapter 22 Rosaceous Shrubs

Miller (1981) and Miller and Kidd (1983) examined (Hitchcock and others 1961). Plants occur singly or in the response of Greene’s mountain ash on seral shrub dense patches interconnected by rhizomes. Stems are fields in northern Idaho to various herbicide treat- slender and yellowish brown to brownish with few ments. Lyon (1966) found that plants were eliminated branches. The simple, alternate, oval to obovate leaves by burning. range from 1 to 2.8 inches (2 to 7 cm) in length; the largest leaves occur near the tips of vegetative and Varieties and Ecotypes flowering branches. The leaves immediately subtending the inflorescence tend to be slightly smaller. Leaves Various selections are used for horticultural are short petiolate, exstipulate, strongly veined, and plantings, but none have been developed for wildlife or coarsely or doubly serrate to shallowly lobulate along wildland uses. the upper one-half to two-thirds of the leaf. Upper leaf surfaces are smooth and shiny; lower surfaces are Sorbus sitchensis ______pale. Inflorescenses are dense, flat-topped terminal Sitka Mountain Ash corymbs 1 to 3.2 inches (3 to 8 cm) in diameter (fig. 29). The tiny, white to pinkish 5-merous flowers are per- Sitka mountain ash grows from 3 to 13 ft (1 to 4 m) fect, perigynous, and regular. Stamens are numerous tall with a rounded crown. It may be distinguished and exceed the petals. Fruits consist of five light readily from Greene’s mountain ash; leaves consist of brown, dehiscent, several-seeded follicles. Flowering 7 to 11 leaflets, each semitruncate to rounded at the occurs from June to early August; fruits ripen from tip and usually at least one-third as broad as long. The mid July to early September. Fruits normally dehisce winter buds, young growth and inflorescences are in October and seeds are shed over time as winter rufous hairy; the calyx is glabrous externally. Stipules are persistent and the fruits are red with a slight bluish cast (Hitchcock and others 1961; Preston 1968). They are dispersed by birds, but some persist on the tree through winter. Sitka mountain ash is distributed from Alaska and the Yukon south to the Cascades in northern Califor- nia and east to British Columbia and northwestern Montana at elevations from 2,000 to 10,000 ft (610 to 3,050 m) (Hitchcock and other 1961). Although it frequently grows in wet soils along streams, it may also be found in forest openings created by natural forest gaps, logging, burning, or other disturbances. Sitka mountain ash produces abundant seed crops. Harvesting, cleaning, storage, germination, and planting requirements are similar to those described for Greene’s mountain ash. Seeds sown late or without adequate wet prechilling do not germinate. Approxi- mately 1 to 5 lb (0.45 to 2.25 kg) of seed may be extracted from 100 lb (45 kg) of fruits (Harris and Stein 1974). Marchant and Sherlock (1984) considered the species a useful high altitude shrub for rocky sites. Man- agement and use of the species are as described for Greene’s mountain ash. It is frequently cultivated as an ornamental.

Spiraea betulifolia ______Bridal Wreath Bridal wreath spiraea, birchleaf spirea, or white spiraea is a low, glabrous, deciduous shrub or sub- Figure 29—Bridal wreath spiraea produces flat shrub ranging from 0.5 to 2 ft (0.2 to 0.6 m) in height corymbs of white flowers.

594 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 22 Rosaceous Shrubs storms dislodge them from the follicles (Hitchcock and leafy softwood cuttings taken in summer. Cuttings others 1961; Stickney 1974c; USDA Forest Service are treated with a root-promoting substance and 1937). rooted under high humidity. Container and 1-0 or 2-0 bareroot seedlings develop rapidly and are easily Ecological Relationships and Distribution handled and transplanted. The genus Spiraea includes about 70 species dis- Uses and Management tributed through temperate and cooler areas of the Northern Hemisphere. About 10 species are found Although it produces abundant available forage, in the western United States, concentrated in mid- bridal wreath spiraea has been rated as worthless for elevations of mountainous areas. They are particu- cattle and poor to fair for sheep (USDA 1937). The larly abundant in the Douglas-fir region of the western dried leaves persist on the plant in fall and may receive United States. The genus is a member of the sub- moderate use after leaves have fallen from other family Spiraeoideae. Chromosome number of the species. Kufeld and others (1973) reported bridal North American species is n = 9. Asian species com- wreath spiraea receives light winter use and heavy monly have chromosome numbers of n = 9 or n = 7 summer and fall use by mule deer. Because of its low (McArthur and Sanderson 1985). palatability, livestock and wildlife rarely damage es- Bridal wreath spiraea occurs from sea level to tablished stands of bridal wreath spiraea. Conse- 11,000 ft (3,300 m) and is distributed from British quently, it may become abundant in heavily grazed Columbia to north-central Oregon and east to Saskatchewan and South Dakota. It grows on a wide sites (USDA Forest Service 1937), spreading vegeta- range of sites from riparian areas to open or wooded tively and from seed. valleys, hills, and rocky slopes. It is particularly abun- Spiraeas, particularly those species that spread dant on logged, burned, or otherwise disturbed mesic rhizomatously, are valuable plants for soil stabiliza- forest sites at intermediate elevations. It is the domi- tion along waterways. Adapted species or varieties nant understory for a number of forest habitat types may be selected for specific areas and conditions. and community types in the Rocky Mountains where Bridal wreath spiraea has also been used to stabilize ponderosa pine, lodgepole pine, grand fir, subalpine roadways, burns, and logged areas. It develops rap- fir, and aspen are the dominant overstory species. idly, producing a spreading root system and rhizomes. Common shrub associates include roses, huckleber- The value of this and other spiraea species for site ries, willows, and snowberries. stabilization is increased by its typically low palatability to game and livestock. Plant Culture A largely unexploited potential of native spiraea is their use as landscape plants. They are easily estab- Bridal wreath spiraea grows in full sun or shaded lished and maintained and produce attractive foliage areas, although most seed is produced on shrubs and showy flowers, although flowering is normally growing in full sunlight. Cleaning involves rubbing restricted to a short period. the follicles to release the seeds and sieving to sepa- Plants developing from seed or transplants mature rate the seeds from the trash. The tiny seeds are rapidly. Those growing in full sunlight may flower and nondormant. Germination reportedly occurs follow- produce seed during the first or usually the second growing season. Although seedlings are reasonably ing 120 days of wet prechilling at 32 to 34 ∞F (0 to 1 ∞C) (Stickney 1974b); this mechanism permits emergence vigorous, their growth may be restricted by competi- shortly after snowmelt. Bridal wreath spiraea may be tion with weeds or seeded grasses. Young seedlings planted by spot seeding disturbed sites in fall or may be trampled by game or girdled by rodents. spring. Seed may be mixed with other shrub seeds, but Most horizontal rhizomes and fibrous roots are lo- should be planted separately from grasses and forbs. cated beneath the mineral soil surface. After fires or Propagating and planting bareroot or container other injury, rhizomes sprout vigorously just behind stock provides for better utilization of the normally the damaged tissue. Roots beneath the surface of limited seed supplies and more rapid development of mineral soil also resprout (Bradley 1984; McLean cover on disturbed sites than does direct seeding. 1967; Fischer and Clayton 1983). Merrill (1982) re- Bareroot stock may be propagated from seed. Hand ported white spiraea achieved a greater volume and seeding may be required in the nursery; it is difficult biomass on burned sites relative to unburned sites to clean the seed to a high level of purity and the light 1 year after burning. This difference was maintained seed does not flow well through a drill. Seed should be over a 4 year study. Rate of recovery was related to covered very lightly and mulched. Bareroot stock may moisture conditions. also be obtained by lining out hardwood cuttings in the spring. Container stock is grown from germinants or Varieties and Ecotypes from hardwood or softwood cuttings (Babb 1959). Hartmann and Kester (1990) recommended the use of None.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 595 Chapter 22 Rosaceous Shrubs

Spiraea densiflora ______Subalpine Spiraea Subalpine spiraea occurs from British Columbia south to northern California, Idaho, and Wyoming at elevations ranging from 2,000 to 11,000 ft (600 to 3,300 m) (Hitchcock and others 1961). It grows along lakes and waterways and on wooded to open rocky slopes, frequently in association with lodgepole pine and western white pine. It is a low, highly branched shrub 1.6 to 3.3 ft (0.5 to 1 m) tall with strong root stocks (fig. 30). Flowers develop in small flat topped Figure 31—Elongate panicles of pink to rose flow- to rounded pink to rose corymbs 0.8 to 1.6 inches (2 to ers characterize Douglas spiraea. 4 cm) wide. Young stems, inflorescences, and lower surfaces of leaves are thickly puberulent. Most material in the Intermountain region belongs to var. splendens. the shrub in winter. Fruits are dry, glabrous, beaked follicles. Seeds are green to brown and less than 0.1 Spiraea douglasii ______inch (2 mm) long. Douglas Spiraea Ecological Relationships and Distribution A relatively tall species, Douglas spiraea grows from Douglas spiraea is distributed from Alaska south to 3 to 8 ft (1 to 2.4 m) in height with erect stems northern California and Montana, occurring prima- branching primarily below the center. The species rily in coastal areas (Davis 1952; USDA Forest Service spreads by rhizomes, often forming thickets in open- 1937). ings along lake and stream margins, bogs, and swamps. It grows on boggy or peaty ground and wet heavy soils Plant Culture subject to annual flooding (Marchant and Sherlock 1984). Stems are pubescent to glabrous. The oblong- Douglas spiraea is easily propagated from cuttings. elliptic leaves are serrate above the center. They are Marchant and Sherlock (1984) obtained 98 percent 1.6 to 4 inches (4 to 10 cm) in length, dark green rooting of softwood cuttings under mist in June and 98 above, and woody beneath. The inflorescence is a percent rooting of hardwood cuttings in fall. They pyramidal rose to pink panicle 1.6 to 11.8 inches (4 to recommended lining out hardwood cuttings in spring 30 cm) long with numerous tiny flowers (fig. 31). Dry and lifting them the same fall for outplanting. Stock flower parts are retained as brown tufts overtopping could also be spring lifted and outplanted. Sprigs or chunks of rhizomes can be dug when dormant and spring planted in areas with moist soils.

Uses and Management Because of their large size, rhizomatous habit, and ease of propagation, both varieties of Douglas spiraea are prime candidates for revegetation of riparian areas, where adapted. They also have potential for use as ornamentals.

Varieties and Ecotypes Spiraea douglasii var. menziesii is separated from var. roseata by its tomentulose inflorescence. It occurs from southern Alaska to northwestern Oregon and northern Idaho. It is one of the most palatable spiraeas and is rated as fair to good fall forage for cattle and Figure 30—Subalpine spiraea inflorescences sheep (USDA Forest Service 1937). The variety roseata are thickly puberulent, flat to rounded corymbs occurs primarily in central Idaho (Hitchcock and oth- of pink to rose flowers. ers 1961). It is a smaller and less palatable shrub.

596 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Stephen B. Monsen Richard Stevens Chapter Nancy L. Shaw 23 Shrubs of Other Families

Introduction ______Numerous genera and species of shrubs occur throughout the Intermountain region in addition to those included in the Asteraceae, Chenopodiaceae, and Rosaceae families. Although shrubs are widespread throughout this region and dominate many areas, species richness is low compared to the shrub flora of the Pacific United States, Chile, western Australia, and South Africa (Stebbins 1975). Generally, evolution proceeds most rapidly when populations are isolated from one another and exposed to different environmental conditions (Dobzhansky 1970; Stebbins 1950); this is the case in the Intermountain region. However, fewer numbers of species within this region are, in part, due to the relatively recent advance and retreat of continental seas, drastic environmental changes, and instability that result in a high rate of extinction (Stebbins 1975). Most species included in this chapter have been used in revegetation programs. Many other shrubs are recognized as important, but information related to their propagation, seed production, germination, or establishment by direct seeding is not available.

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Chapter Contents by Species

Acer glabrum (Rocky Mountain maple) ...... 600 Rhus aromatica (Skunkbush sumac)...... 608 Rhus glabra (Rocky Mountain smooth sumac) ...... 611 Mahonia aquifolium (Shining barberry) ...... 613 Mahonia fremontii (Fremont barberry)...... 614 Mahonia repens (Oregon grape, creeping barberry, mountain holly, creeping mahonia) ...... 614 Alnus incana (Thinleaf alder) ...... 616 Betula glandulosa (Bog birch)...... 618 Betula occidentalis (Water birch) ...... 620 Betula papyrifera (Paper birch) ...... 622 Lonicera ciliosa (Orange honeysuckle) ...... 623 Lonicera involucrata (Bearberry honeysuckle) ...... 623 Lonicera utahensis (Utah honeysuckle) ...... 624 Sambucus cerulea (Blue elderberry)...... 624 Sambucus racemosa (Red elderberry) ...... 625 Symphoricarpos albus (Common or white snowberry) ...... 627 Symphoricarpos longiflorus (Desert snowberry) ...... 627 Symphoricarpos occidentalis (Western snowberry, wolfberry) ...... 628 Symphoricarpos oreophilus (Mountain snowberry) ...... 629 Cornus stolonifera (Redosier dogwood)...... 630 Cupressus arizonica (Arizona cypress) ...... 632 Juniperus horizontalis (Creeping juniper) ...... 633 Juniperus osteosperma (Utah juniper) ...... 635 Juniperus scopulorum (Rocky Mountain juniper) ...... 638 Elaeagnus commutata (Silverberry) ...... 640 Shepherdia argenta (Silver buffaloberry)...... 641 Shepherdia canadensis (Russet buffaloberry, soapberry) ...... 642 Shepherdia rotundifolia (Roundleaf buffaloberry) ...... 643 Ephedra nevadensis (Nevada ephedra) ...... 644 Ephedra viridis (Green ephedra) ...... 645 Arctostaphylos uva-ursi (Bearberry manzanita) ...... 646 Quercus gambelii (Gambel oak) ...... 648 Fraxinus anomala (Singleleaf ash) ...... 650 Eriogonum heracleoides (Wyeth or whorled eriogonum, buckwheat)...... 652 Eriogonum umbellatum (Sulfur eriogonum, buckwheat) ...... 653 Eriogonum wrightii (Wright eriogonum, buckwheat) ...... 654 Clematis ligusticifolia (Western virginsbower)...... 654 Ceanothus species ...... 656 Ceanothus cuneatus (Wedgeleaf ceanothus) ...... 656 Ceanothus greggii (Desert ceanothus) ...... 657 Ceanothus integerrimus (Deerbrush ceanothus)...... 658 Ceanothus lemmonii (Lemmon ceanothus) ...... 659 Ceanothus prostratus (Prostrate ceanothus) ...... 659 Ceanothus martinii (Martin ceanothus) ...... 661 Ceanothus sanguineus (Redstem ceanothus) ...... 662 Ceanothus velutinus (Snowbrush ceanothus) ...... 665 Salix species (Willow) ...... 668 Salix bebbiana (Bebb willow, beaked willow) ...... 670 Salix boothii (Booth willow) ...... 672

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Salix drummondiana (Drummond willow, beautiful willow, blue willow) ...... 673 Salix exigua (Coyote willow, dusky willow, narrowleaf willow, sandbar willow) ...... 675 Salix geyeriana (Geyer willow) ...... 676 Salix glauca (Grayleaf willow, glaucous willow) ...... 677 Salix lasiandra var. caudata (Whiplash willow) ...... 679 Salix lasiolepis (Arroyo willow) ...... 680 Salix lutea (Yellow willow, shining willow) ...... 681 Salix planifolia (Plainleaf willow) ...... 683 Salix scouleriana (Scouler willow, fire willow, black willow, mountain willow, nuttall willow) ... 684 Salix wolfii (Wolf willow) ...... 686 Philadelphus lewisii (Lewis mockorange) ...... 687 Ribes aureum (Golden currant) ...... 689 Ribes cereum (Wax currant) ...... 691 Ribes viscosissimum (Sticky currant) ...... 693 Penstemon fruticosus (Bush penstemon) ...... 694 Celtis reticulata (Netleaf hackberry) ...... 696

Consequently, they are not widely used in large-scale Shrubs discussed in this chapter occupy rather spe- seedings where uniform stands are expected. A num- cific sites. Mountain snowberry, Rocky Mountain ber of important species could be more widely utilized maple, and Gambel oak are confined to specific in artificial revegetation and increased through man- elevations and climatic conditions, yet occupy rather agement if more information related to their culture large acreages, often as the dominant species. Other was available. shrubs, including green ephedra and singleleaf ash, Sixty-seven species representing 20 families are are common throughout the southern portion of the discussed in this chapter. These species occur over a Intermountain area but do not extend into the colder wide range of conditions and habitats. Certain species northern section. By contrast, red elderberry and dwarf are common and widespread; others are less abundant snowberry occupy only mountainous situations. and often restricted to localized conditions. However, A characteristic of many shrubs discussed in this all are important. In combination, these shrubs offer a chapter is their close association or relationship to soil diverse range of material for planting and manage- or site conditions. Rather extensive areas of willow ment. Eriogonum and Ceanothus are examples of two occur through many vegetative communities but are widespread genera discussed in this chapter. Approxi- most often confined to moist soils. The relationship of mately 130 species of Eriogonum, including shrubs blue elderberry, bush penstemon, and other shrubs and broadleaf herbs, occur within the Intermountain that are also adapted to specific soil conditions is less region (Welsh and others 1987). Of these, fewer than well understood. 10 have been evaluated or used for range or wildland Introduced grasses have not been found to be as plantings (Miles and Meikle 1984; Plummer and oth- specifically adapted as many of these shrubs. This is ers 1968; Sampson and Jespersen 1963). By contrast, an important consideration; some shrub-dominated about two dozen species of Ceanothus are reported in associations have been planted to introduced species, this same region and slightly more than half have been mostly grasses. Attempts have been made to substi- studied for wildlife, watershed, or horticultural uses. tute other species for native shrubs. Many substitute In most instances only one or two species of any genus plants do not provide the forage or cover values ex- discussed in this chapter have been studied as poten- pected or simply are not adapted for long-term persis- tial revegetation species. Seed or planting stock of tence on some sites. Thus, it is important to learn most species is often relatively scarce. restoration and management practices required to perpetuate endemic shrub associations.

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Some shrubs are particularly important during aesthetic and cultural resources that are unique to the various stages of succession. Species of redstem Western United States. ceanothus, snowbrush ceanothus, and red elderberry Important characteristics of a number of shrubs recover well following fires and assume dominance are listed in table 1. Seeding recommendations for after coniferous forests have burned. These shrubs major vegetative types and conditions are discussed provide important habitat, but unless they are able to in chapter 17. Shrubs adapted to these situations are perpetuate during cyclic fire events, game and live- included in the seeding recommendations. Seed char- stock habitat may eventually diminish as conifers acteristics are found in chapters 24 through 27. gain dominance. Many shrubs occur as components of associated forest and herbaceous communities. Considerable fluc- Family Aceraceae ______tuation in species composition often occurs as a result Acer glabrum of fires, and from extremes in climatic events, grazing, Rocky Mountain maple and disease. The opportunity to reestablish plants on these disturbances is, in part, dependent upon use of Description—Rocky Mountain maple is a long- ecotypes with appropriate adaptive features. In an lived, shade-tolerant, deciduous shrub or small tree attempt to enhance the shrub component where desir- native to North America (Van Dersal 1938). Plants able species have been lost, planting in some situa- vary in height from 6 to 20 ft (1.8 to 6.1 m) (Harrington tions, has been very successful. Characteristics that 1964), and also demonstrate considerable variation in ensure the survival of shrubs under adverse condi- stature, leaf form, and fruit shape (Kartesz and Kartesz tions are important in selecting plants to rehabilitate 1980). Some plants reach 20 to 30 ft (6.1 to 9.1 m) in mine disturbances and other harsh sites. height with multiple trunks up to 1 ft (0.3 m) in Some shrubs are often the only or dominant species diameter, but they occur most often as shrubby plants to occur on harsh and disturbed sites. Soils throughout with multiple stems less than 15 ft (4.6 m) tall many areas of the Intermountain region are infertile (Haeussler and Coates 1986; Hitchcock and Cronquist and do not support large numbers of species. It is 1973). Leaves are simple, opposite (Alexander and unlikely that successional changes will result in the others 1987), palmately three- to five-lobed or three replacement of many shrubs with other, equally foliate, and mainly 0.8 to 3.2 inches (2 to 8 cm) wide adapted species in such areas. Certain species of (Welsh and others 1987) (fig. 1). The herbage is glabrous. buckwheat and penstemon are able to populate harsh, exposed sites. Many of these sites are important midwintering areas where high quality forage is essential for big game animals. In addition, these exposed sites are often invaded by weeds that may enter and, subsequently, spread to adjoining locations. With- out the presence of adapted species, weedy plants can invade open and seemingly harsh situations. Many shrubs are important forage plants. Some are browsed readily throughout the year; others are selectively grazed during certain seasons. For example, Gambel oak, Rocky Mountain maple, redstem ceanothus, and Martin ceanothus are often heavily browsed and can contribute a considerable amount of forage throughout the entire year. Green ephedra, Utah juniper, and Oregon grape usually are grazed during specific seasons. Woody plants provide not only forage, but are equally important as cover and concealment for wildlife. Utah juniper, Gambel oak, skunkbush sumac, Rocky Moun- tain maple, and willows are very important for this purpose. Many shrubs discussed in this chapter are also useful for soil stabilization, watershed protection, and ground cover. Deerbrush ceanothus, Scouler willow, mountain snowberry, thinleaf alder, and golden currant are examples of species useful for conservation plantings. The native shrub communities provide Figure 1—Leaves of Rocky Mountain maple are opposite and palmately five-lobed.

600 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

(con.)

A,PP,MB,JP,R,MS

A,WM,PP,MB,R MB,JP,BS,WS,BB MB,JP,BS,WS A,PP,MB,JP,MS,BS A,PP

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Disease and insect resistance insect and Disease

Flooding tolerance Flooding

Shade tolerance Shade

Compatibility with other species other with Compatibility

Soil stability Soil

Seed production Seed

Grazing tolerance Grazing

Edible foliage retained fall and winter and fall retained foliage Edible

Palatability, summer growth summer Palatability,

Palatability, early spring growth spring early Palatability,

Availability of current growth current of Availability

Herbage yield Herbage

Growth rate Growth

Natural spread Natural

Persistence

Final establishment Final

Initial establishment Initial

Germination

Ease of transplanting of Ease Ease of seeding of Ease

25445444543424355545 33423432343445244234 32223522342445244124 44223442223354234434 44213432242244254424 23334444433434244443SA,A,PP,WM,R 25444445432424344443 25444445422424344443 55545444443445545424 53543333443345434224 25224554442324154543 52411431333435133125 54333434443434344444 42223322333344143124 44224433343444243534 44334544443434344434 55444445553435354334 53323333433445243214 44335543433435254424 54433542223445253424 55445555553435344434 55445555553445354434 55434444553445354324 45444414344424444433 54224533445435354324 34324544545435454433SA, 34224533344434244444 43433443443455435124 55444544455524344443 34323323444434334434 Ease of cleaning seed cleaning of Ease

4 4

3 3 3 3 5 5 2 3 3 3 3

the species is adapted is species the

to which to —Species characteristics of selected Intermountain shrubs. types Vegetative a

Table 1 Alder, thinleaf 3 Apache plume 1 Ash, singleleaf 4 Barberry, creeping Bearberry 3 Birch, bog 3 Birch, paper 3 Birch, water 3 Bitterbrush, antelope Bitterbrush, desert Blackcap 3 Blackbrush 3 Buckthorn, cascara Buffaloberry, roundleaf Buffaloberry, russet Buffaloberry, silver Ceanothus, deerbrush Ceanothus, Fendler Ceanothus, Martin Ceanothus, prostrate Ceanothus, redstem Ceanothus, snowbrush Ceanothus, wedgeleaf Cherry, Bessey 3 Cherry, bitter 3 Chokecherry, black Cinquefoil, bush 4 Cliffrose, Stansbury Currant, golden 3 Currant, sticky 3

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 601 Chapter 23 Shrubs of Other Families

(con.)

BS,R

,BS,WS

A,PP,MB,JP,BS MB,JP,MS,BS A,PP,WM,MB,JP,R PP,MB,MS,WM

A,MB,JP,MS SA,A,PP,MB MB,JP,MS,BS,WS,SS,BB MB,JP,BS,WS,SS,BB

SA,A,PP,MB,JP,MS,BS,WS MB,JP,BS,WS BS,WS,SS,BG,IS WM,PP,MB,MS,R WM,PP,MB,MS,R PP,MB,JP,MS SA,A,WM,MB A,PP A,PP,MB JP,BS,WS,SS,BB JP,BS,WS,SS,BB A,PP A,MB,JP,MS,BS

MB,JP,MS,BS,WS,SS,BB

PP,MB,JP,MS,WS

Disease and insect resistance insect and Disease

Flooding tolerance Flooding

Shade tolerance Shade

Compatibility with other species other with Compatibility

Soil stability Soil

Seed production Seed

Grazing tolerance Grazing

Edible foliage retained fall and winter and fall retained foliage Edible

Palatability, summer growth summer Palatability,

Palatability, early spring growth spring early Palatability,

Availability of current growth current of Availability

Herbage yield Herbage

Growth rate Growth

Natural spread Natural

Persistence

Final establishment Final

Initial establishment Initial

Germination

Ease of transplanting of Ease Ease of seeding of Ease

44224433344434234334 43334413333544333234 45334445444434345444 55334534444434344434 44125535545525345335A,PP,MB,JP,MS, 25542234224414433244 44125535545525344445 52543422344555334115 52543422344455234115 53445354334444333323PP,MB,JP,MS

53445444344444334424 44324524232534434224 43414523333525334155 44224532433434244334 34124552342334234343 45334544332422333322 35214533445544244434 45424534444424244435 45324544443344253324 44442433334535343135 42412512225115233135 53333533445425343323 53114522242255242324 53214452343255332334 51115552232355214224 Ease of cleaning seed cleaning of Ease

c c c

2 3 3

5 3

3 2 3 3 3

3 3 3 3

the species is adapted is species the

to which to Vegetative types Vegetative

(Con.) a

Table 1

Currant, wax 3 Cypress, Arizona Dogwood, redosier Elaeagnus, autumn Elderberry, blue 3 Elder, box 3 Elderberry, red 3 Ephedra, green 5 Ephedra, Nevada Eriogonum, sulfur

Eriogonum, Wyeth Forestiera, New Mexico Greasewood, black Hackberry, netleaf Hawthorn, Douglas Honeylocust 3 Honeysuckle, bearberry Honeysuckle, Utah Honeysuckle, orange Hopsage, spineless Hopsage, spiny 3 Indian apple 3 Juniper, Rocky Mountain Juniper, creepingJuniper, Utah 5 5

602 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

(con.)

A,PP,WM,MB

A,WM,PP,MS

M,MB,R

B,JP,MS,BS,WS,BB

,PP,MB,JP,MS

WS,SS,BG MB,PJ,MS,BS,WS, SS,BB, C,AW PP,MB,JP,MS A,MB,JP A,PP,MB

A,WM,PP,MB

PP,MB,JP

A,MB,JP A,WM,PP,MB,MS,R MB,JP,BS,WS,SS,BB A,PP,MB,MS

A,PP,MB,JP,MS, BS,WS,SS

A,MB,JP,MS,BS,WS, SS,BG,BB,IS SA,A,MB,SS SA,A,PP,MB,JP, MS,BS,WS,SS, BG,IS JP,MS,BS,WS,SS BS,WS,SS,BG,BB

SA,A,PP,MB,JP,MS

Disease and insect resistance insect and Disease

Flooding tolerance Flooding

Shade tolerance Shade

Compatibility with other species other with Compatibility

Soil stability Soil

Seed production Seed

Grazing tolerance Grazing

Edible foliage retained fall and winter and fall retained foliage Edible

Palatability, summer growth summer Palatability,

Palatability, early spring growth spring early Palatability,

Availability of current growth current of Availability

Herbage yield Herbage

Growth rate Growth

Natural spread Natural

Persistence

Final establishment Final

Initial establishment Initial

Germination

Ease of transplanting of Ease Ease of seeding of Ease

43324532224545244134 44424543444545444224 34344544332433333323 33422333445423444434 22212333234423335533 23433444443244332232A,PP,WM,MB,MS 43323442333234322233 32222433434234224244SA,

43433433333554333313

43432433343444342213 43443444344424343323A 44433443444244234334A,WM,PP,MB,MS 21432423441525353434 44334433342255233343 44544443335414333133 55444433544433323431 23324443334414343344W

24334453333434334224 25444553234445354114M 24334544344453442344 25434443534445344224 24444455553355454223

25444554333445343115 25445444445344353234 43234423442354243243SA, 45234534434425244444 Ease of cleaning seed cleaning of Ease

d d d d d

d d

3 1 3

2 3

4 5 4

4 4

the species is adapted is species the

to which to Vegetative types Vegetative

(Con.) a

Table 1

Kochia, gray-molly Kochia, forage 3 Locust, black 3 Maple, bigtooth 3 Maple, Rocky Mountain Mockorange, Lewis mountain ash, Greene’s Mountain lover 2 Mountain mahogany, curlleaf 2 Mountain mahogany, littleleaf 2 Mountain mahogany, true Ninebark, mallowleaf Oak, Gambel 5 Oceanspray, creambush Peachbrush, desert Penstemon, bush Plum, American 3 Rabbitbrush, Douglas

Rabbitbrush, dwarf

Rabbitbrush, Parry Rabbitbrush, rubber

Rabbitbrush, small Rabbitbrush, low 4 Rabbitbrush, spreading Rockspirea 4 Rose, Woods 3

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 603 Chapter 23 Shrubs of Other Families

(con.)

A,WM,PP,MB

JP,BS,SS,BG,BB,C,AW

JP,BS,WS,SS,BG,BB MB,JP,MS,BS,SS

BB,WS,SS MB,MS MB,JP,MS,BS,SS JP,BS,WS,SS,BG

WS,SS,BB

MS,BS,WS MB,MS,BS,WS,JP,MS JP,BS,WS,SS,BG,BB,C,AW

WS,SS,BG MB,JP,MS,BS,WS,SS, BG,BB,IS MB,JP,BS,WS,SS,BG,IS WS,SS,BG,BB WS,SS,BG,BB JP,BS,WS,SS,BG,BB

JP,BS,WA,BB SA,A,PP,MB,JP,MS PP,MB,JP,MS,BS,BB WM,PP,MB

A,JP,BS,WS,BB SA,A,PP,MB,JP,MS WM,MB,BS

PP,MB,JP,MS PP,MB,JP,MS,BS,BB

Disease and insect resistance insect and Disease

Flooding tolerance Flooding

Shade tolerance Shade

Compatibility with other species other with Compatibility

Soil stability Soil

Seed production Seed

Grazing tolerance Grazing

Edible foliage retained fall and winter and fall retained foliage Edible

Palatability, summer growth summer Palatability,

Palatability, early spring growth spring early Palatability,

Availability of current growth current of Availability

Herbage yield Herbage

Growth rate Growth

Natural spread Natural

Persistence

Final establishment Final

Initial establishment Initial

Germination

Ease of transplanting of Ease Ease of seeding of Ease

35545555553354443233 35544454343344333233 45545554333554443114 22213322225125224124 33444443333345232144 34344443332544334444 34545554332235232244 35445555553455344333SA, 22322222223334234234 45545554553455443344SA,A,MB,MS,WS 23323332233435233224 35545555553354444334 35545444443354343223

43433434445543321343 54443435543555545122 44333343344545343133 44323432445553211343 43223454445423431343 42212332333535244123

35445444333344344224 53333433344434344423 53334433344434344324 45334423342334434344 55334334444424243444MB 55233434445324144224 55334544445425255444 55334434344424244434 34435534452254244343A,WM,PP,MB,MS

55224434443434444224 55224533443435343234 Ease of cleaning seed cleaning of Ease

d d d d d d d d d d d d d

4 4

4 5

4 4

3 3

3 3 3 3 3

the species is adapted is species the

to which to Vegetative types Vegetative

(Con.) a

Table 1

Sagebrush, basin big Sagebrush, Bigelow

Sagebrush, black Sagebrush, bud 4 Sagebrush, early Sagebrush, fringed Sagebrush, low 4 Sagebrush, mountain big

Sagebrush, pygmy Sagebrush, silver Sagebrush, stiff 4

Sagebrush, threetip Sagebrush, Wyoming big Saltbush, Castle Valley clover 4 Saltbush, fourwing Saltbush, Gardner Saltbush, mat 5 Saltbush, quailbush Saltbush, shadscale Sandsage 4

Serviceberry, Saskatoon Serviceberry, Utah Silverberry 3 Snowberry, common Snowberry, longflower Snowberry, mountain Snowberry, western Spiraea, birchleaf Sumac, Rocky Mountain smooth 2 Sumac, skunkbush

604 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

WM,R

WM,PP,R

A,WM,PP,MB

A,PP,MB,JP,MS,BS,BB A,WM,PP,R WM,A,PP,IS,R

A,WM,PP,MB,R

A,PP,P-J,WM,R MB,JP,BS,WS,SS,BG,BB,IS

SA,A,PP,MB,MS

Disease and insect resistance insect and Disease

Flooding tolerance Flooding

Shade tolerance Shade

Compatibility with other species other with Compatibility

Soil stability Soil

Seed production Seed

Grazing tolerance Grazing

Edible foliage retained fall and winter and fall retained foliage Edible Palatability, summer growth summer Palatability,

= Shadscale saltbush; WM = Wet and semiwet meadows; BG = Black greasewood; PP = Ponderosa pine; BB = Blackbrush;

= Basin big sagebrush; AW = Annual weeds. Palatability, early spring growth spring early Palatability,

Availability of current growth current of Availability

Herbage yield Herbage

Growth rate Growth

Natural spread Natural

Persistence

Final establishment Final

Initial establishment Initial

Germination

Ease of transplanting of Ease Ease of seeding of Ease

25224553342324154543 24323443334414244324 24424455455354344445 24424445553344534334 24424455455354324445SA,A,PP, 25424425454444454444 25424445554444454434A, 25424445554444454434A, 25424445454444344444 25544445334445335124

35114235333534154215 Ease of cleaning seed cleaning of Ease

2 2

the species is adapted is species the

to which to Vegetative types Vegetative

(Con.) a

Vegetative type or community to which the species is adapted: SA = Subalpine; WS = Wyoming big sagebrush; A = Aspen-conifer; SS Key to ratings: l = PoorÐdifficult; 2 = Fair; 3 = Medium; 4 = Good; 5 = ExcellentÐeasy.

Less than 15 percent purity.

In the flesh.

a b c d

Table 1

Thimbleberry 3 Virginsbower, western Willow, Booth 2 Willow, coyote 2 Willow, Drummond Willow, purpleosier Willow, Scouler 2 Willow, whiplash 2 Winterfat, common MB = Mountain brush; IS = Inland saltgrass; JP = Juniper-pinyon; R = Riparian; MS = Mountain big sagebrush; C = Cheatgrass; BS Wormwood, oldman

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 605 Chapter 23 Shrubs of Other Families

Twigs are usually gray with smooth, reddish-brown soil, with high concentrations of organic matter in the bark. Flowers are monoecious or dioecious, appearing upper portions of the soil profile (Krajina and others in loose, racemose corymbs (Harrington 1964). Fruits 1982; Mueggler 1965). Plants also occur on rocky and consist of two fused samaras about 1 to 1.5 inches (2.5 gravelly soils and on steep slopes with shallow soils, to 3.8 cm) long, with widely divergent wings, each where a considerable amount of litter has accumu- containing a single seed (Harrington 1964; Hitchcock lated. At both upper and lower elevation limits, the and Cronquist 1973; Olson and Gabriel 1974). distribution of Rocky Mountain maple is influenced by topographic features and aspects. Ecological Relationships and Distribution— Rocky Mountain maple usually grows on sites that Rocky Mountain maple occurs from Alaska south to receive between 16 and 30 inches (400 and 760 mm) of California, Arizona, New Mexico, and Nebraska (Little annual precipitation. Mature stands will usually per- 1979; Welsh and others 1987). In the Pacific North- sist with limited dieoff during periods of extended west, Rocky Mountain maple is primarily a coastal drought. A decrease in herbage production may occur species, but it is distributed eastward into Idaho, if the precipitation level falls below normal, but stand Montana, and western Colorado at elevations between density usually remains unchanged. 5,000 and 10,500 ft (1,500 and 3,200 m) (Hitchcock and Rocky Mountain maple may occur in dense, re- Cronquist 1973; Harrington 1964). It occurs at eleva- stricted patches or clumps; yet, open diverse stands tions of 5,000 to 9,000 ft (1,500 to 2,700 m) in conifer often occupy an entire slope or small drainage (fig. 2). forests of the Sierra Nevada Mountains, northern When found on south or west aspects, it may be Coast Ranges, and into western Nevada (Munz and confined to small patches surrounded by bunchgrasses Keck 1959). Welsh and others (1987) reported the plant or more xeric species. Mountain brome and slender occurs in all Utah counties at elevations between wheatgrass frequently occur as an understory with 5,440 and 10,300 ft (1,700 and 3,100 m). this species. These two grasses may increase or decline Rocky Mountain maple is frequently a seral species, dramatically from year to year, but the density of but it is able to persist as an understory in climax overstory maple remains quite consistent. conifer stands due to its shade tolerance (Mueggler Grazing, wildfires, and other impacts have not seri- 1965; Ryan and Noste 1985). It occurs intermixed with ously reduced the distribution or density of Rocky Douglas-fir, lodgepole pine, ponderosa pine, spruce- Mountain maple. Cutting and burning temporarily fir, and quaking aspen. However, it does not occur as reduce some stands, but natural recovery is quite a common understory species with aspen in Utah rapid. Weedy species have not seriously affected the (Mueggler 1988). It occurs along streambanks in moun- presence or vigor of Rocky Mountain maple even tainous areas, in moist canyon bottoms, and on slopes though heavy grazing has removed the native under- (Mozingo 1987; Wasser 1982), primarily in moist habi- story in some areas. tats (Lotan 1986). This species is frequently associated with mallowleaf ninebark, Saskatoon serviceberry, Scouler willow, Greene’s mountain ash, chokecherry, mountain lover, and Utah honeysuckle (Atzet 1979; Johnson and Simon 1987; Mauk and Henderson 1984; Steele and others 1981; USDA For- est Service 1973; Wasser 1982). Within the Intermountain region, Rocky Mountain maple occurs primarily in mountain brush communities. The species exists over a wide range of sites from well-drained to moist situations. Plants grow intermixed with Gambel oak, chokecherry, and Woods rose in mesic, brushland sites. At lower elevations, its range overlaps with pinyon and juniper, although it usually does not grow in mixtures with these conifers. Where pinyon or juniper occur with maple, the conifers are usually restricted to the more arid sites. In arid regions it is restricted to streams and moist canyons (Mozingo 1987; Wasser 1982), existing as an important species in riparian communities (Brown and others 1977; Mozingo 1987; Myers 1987). It occupies both warm and cool wet areas. Figure 2—Rocky Mountain maple may occur as In some situations, Rocky Mountain maple appears individual shrubs, or more often in small thickets in to be dependent on the presence of a well-developed mountain brush communities.

606 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Rocky Mountain maple is particularly important in To date, most seedings have been confined to sites many of the conifer forest communities. It is not seriously where the species previously existed. When seeded diminished by a closing canopy (Antos 1977; Habeck “offsite,” weak and erratic stands have developed. 1970; Stickney 1985). It can recover immediately follow- Plants appear to be quite dependent on soil conditions ing fire (Cholewa and Johnson 1983; Mueggler 1965) or to survive and flourish. When seeded or transplanted logging due to its resprouting capabilities. Recovery is on mine wastes, plants have done poorly. However, if from stem resprouting, not by root suckering or rhizome sites are top soiled, better stands have established and formation (Haeussler and Coates 1986). persisted. Natural seedlings are usually not abundant. When Plant Culture—Seeds usually mature in August or found, they are encountered in shady areas with little September (Olson and Gabriel 1974). Some stands competition (Steele and Geier-Hayes 1992). Under produce seeds once every 2 to 3 years (Olson and these conditions seedlings have been reported to grow Gabriel 1974). However, individual plants scattered slowly. throughout a stand usually produce some seeds each Rocky Mountain maple is easily cultured and has year. In addition, certain plants have been observed to been planted in commercial landscape programs. produce a viable crop of seeds every year for 3- to 5- Transplants or wildlings are used in conjunction with year periods. Individual plants often produce an ex- other species in formal plantings. traordinary amount of seeds. Seeds also tend to ripen Transplanting 1- to 3-year-old nursery stock has more uniformly on certain plants; consequently, seed improved stand establishment in areas with poor collection is usually confined to a few plants. Seeds soils or where some understory competition exists. often ripen over a period of 1 month. Older transplants are able to establish and begin Seeds usually remain on the plants for 2 to 4 weeks growth quite rapidly without a prolonged period of before being dislodged by wind. When mature, the adjustment. fruits turn from green to yellow or reddish brown. Seeds can be collected by hand stripping or flailing the Uses and Management—Rocky Mountain maple bush. Green or immature seed should not be collected. has been planted to provide diversity and enhance Seeds should be inspected for insect damage and wildlife habitat. Though it is often confined to small immature embryos before collection. Some collections areas, it provides valuable thickets for protection and exhibit about 50 percent germination (Swingle 1939). concealment. It is a useful forage species and occurs in This percentage can be improved, however, if care is important winter habitats for big game animals (Arno taken to collect mature, well-developed seeds. There and Hammerly 1977; Beetle 1962; Gaffney 1941; Keay are about 13,430 seeds per lb (29,600 per kg) (Olson 1977; Martinka 1976; Young and Robinette 1939). and Gabriel 1974). Rocky Mountain maple normally supports a produc- The wings are usually detached from the fruits to aid tive complex of understory herbs. These sites are in handling and planting. Once the wings are re- important areas for summer grazing by big game and moved, empty seeds can be removed by flotation. This livestock. Stands of Rocky Mountain maple that re- is advisable to permit regulation of seeding rates. cover following burning provide considerable forage Seeds should be dried to a water content of 10 to 15 for big game, livestock, and nongame animals. In most percent before storage (Olson and Gabriel 1974). Seeds situations plants may grow too tall to be reached by can be stored for 1 to 2 years in sealed containers at 35 animals, yet they provide excellent cover (Johnson to 41 ∞F (1.7 to 5 ∞C) without serious loss of viability and Simon 1987; Larson and Moir 1987). In more arid (MacArthur and Fraser 1963). Seeds have dormant situations, the shorter growing forms remain avail- embryos that require 6 months of wet prechilling to able to browsing animals. Rocky Mountain maple is a release dormancy (Shaw 1984). Consequently, fall palatable species, particularly to big game (Beetle seeding is advised. However, wet prechilled seeds can 1962; Keay 1977; Martinka 1976; Mitchell 1950; Smith be spring planted, particularly under nursery condi- 1953), but is also used by other animals, including tions. Seeds dispersed on the soil surface do not main- ruffed and blue grouse (Steele and others 1981) and tain viability (Kramer 1984). small mammals (Martin and others 1951). Cleaned seeds can be planted with most conven- Rocky Mountain maple grows in moist areas. When tional seeders, but most plantings have been restricted disturbed, these sites can produce sediment and in- to small, carefully prepared areas where hand seeding crease runoff. These sites are important for watershed has been employed. Seeds should be planted at a depth stability and may require immediate attention if dis- of 0.5 to 1.5 inches (1.3 to 3.8 cm). Broadcast planting turbed. Rocky Mountain maple is able to recover without any followup methods for covering the seeds is following disturbances and can provide soil and not advised. Planting weedy sites should be avoided. streambank protection. Seedlings have grown well and attained considerable Both Rocky Mountain maple and bigtooth maple are size in 1 to 2 years. attractive plants, particularly during the fall when

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 607 Chapter 23 Shrubs of Other Families leaves turn red and yellow. Certain plants and clumps and others 1987); some may reach heights of 15 ft exhibit early fall coloration, others are particularly (4.6 m) (Thornburg 1982). Crown diameter often ex- brilliant in color, and some may retain their leaves for ceeds the height and may be more than 30 ft (9.1 m) extended periods. Also, considerable differences in (Plummer and others 1968; USDA Forest Service growth habit and mature size occur among ecotypes. 1976c). Stems are numerous, woody, spreading, highly Many native stands have high aesthetic appeal and branched, and brown hairy when young; they develop are important viewing areas and recreational sites. a gray bloom with age (Barkley 1937; Wasser 1982). These areas are frequently managed for scenic use. Leaves are 1 to 3 inches (3 to 7 cm) long, trifoliate, Rocky Mountain maple has not been used as exten- petiolate, and puberulent beneath. The terminal leaf- sively in windbreaks and conservation plantings as let is fan shaped, shallowly to deeply incised or lobed, other maples. and coarsely crenate. Lower leaflets are half as large and shallowly crenate. The resinous, acrid sap of the Varieties and Ecotypes—None. twigs and leaves is nonpoisonous, but produces a disagreeable odor, hence the common name, skunkbush Family Anacardiaceae______sumac (Stubbendieck and others 1986). Plants are polygamo-dioecious. The inflorescence consists of a Rhus aromatica cluster of spikes located near the branch apex (Goodrich Skunkbush sumac and Neese 1986). Buds and flowers are inconspicuous, pale yellow, and mostly imperfect. Fruits are orange Description—Skunkbush sumac is an unusually red, sticky, berrylike drupes, each containing a single persistent and highly variable species (fig. 3). Welsh bony seed (Brinkman 1974g; Stubbendieck and others and others (1987) described two varieties of this spe- 1986; Welsh and others 1987). cies complex that occur in Utah. Rhus aromatica var. simplicifolia has simple leaves and occupies xeric Ecological Relationships and Distribution— sites in southeastern and southwestern Utah. Rhus Skunkbush sumac is widespread and highly variable; it aromatica var. trilobata leaves are trifoliolate, and ranges from Alberta south to Iowa and Mexico, and plants grow on mesic sites including streambanks, west to southeastern Oregon and California (Harrington seeps, and riparian communities. Shrubs occur singly 1964). It is found at elevations from 3,500 to 8,000 ft or in dense patches and may be connected by woody (1,100 to 2,400 m) (Hitchcock and others 1961; USDA rhizomes, which often exceed 20 ft (6.1 m) in length Forest Service 1937) on sites with annual precipita- (Sanford 1970). Root systems are deep and extensively tion between 10 and 20 inches (250 and 510 mm) branched. Plants are deciduous, woody, spreading, (Wasser 1982). It grows in the mountain brush and and moundlike to erect, often forming clumps or pinyon-juniper types of the Intermountain region and thickets. They normally range from 2 to 7 ft (0.6 to the plains and foothills of eastern Wyoming and Mon- 2.1 m) in height (Hitchcock and others 1961; Welsh tana (Welsh and others 1987). It is a primary or secondary species growing in sand hills of the plains and Southwest (Wasser 1982). At lower elevations it may be found with blackbrush, basin big sagebrush, or Wyoming big sagebrush. It also occurs in openings in oakbrush and ponderosa pine forests (Vories 1981). Plants growing in rocky foothills are usually smaller in stature and less productive than the more typical upright forms of the mountain brush communities (Welsh and others 1987). Skunkbush sumac is generally drought and cold tolerant, but these characteristics vary among ecotypes. Skunkbush sumac is commonly found along stream courses where it occurs intermixed with other shrubs or as an understory with cottonwoods. Although dense stands are usually not encountered, scattered plants or thickets may be widely distributed in openings along riparian areas; plants require full sunlight to partial shade. On dry sites it occurs in swales and drainageways where it may receive extra moisture. It Figure 3—Skunkbush sumac established from a requires well-drained soils with deep water tables direct seeding provides important wildlife habitat. (Wasser 1982), and although it tolerates short-term

608 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families flooding where it occurs along streams, it is not ca- release embryo dormancy (Babb 1959; Heit 1970; pable of surviving long-term flooding. Swingle 1939). Duration of acid treatment and wet Skunkbush sumac occurs on a wide range of soils prechilling periods required vary among seedlots. ranging from sandy textured soils to heavy clay-tex- Physical rupturing of the seedcoat using mechanical tured materials (Johnson and others 1966). In Utah, it methods also increases germination (McKeever 1938), is particularly well adapted to heavy shale outcrops, but procedures have not been developed to treat large and sometimes salty outcrops, where few other shrubs seedlots. Fall seeding often provides the necessary are adapted (Plummer 1977). It grows on both acidic cold treatment to release embryo dormancy (Brinkman and basic soils including slightly alkaline soils (Johnson 1974g), but very erratic results have occurred, possi- and others 1966; Vories 1981). It occurs on deep fertile bly due to differences in seedcoat imposed dormancy soils and also on shallow, rocky, less-developed soils. (Boyd 1943; Monsen 1987; Monsen and McArthur 1985). Plant Culture—Fruits of skunkbush sumac form Erratic seed germination is the principal factor dense clusters near the ends of small branches. The preventing extensive seeding of this shrub. Differ- clusters are scattered over the surface of the entire ences in germinability are not entirely related to bush. Flowers appear from March to April (Brinkman ecotypes or site of collection. Plants from sites in 1974g). Fruits ripen from June to September (Plummer north-central Wyoming and south-central Montana and others 1968; Van Dersal 1938), and many remain are in considerable demand for mine restoration, but on the shrub until midwinter. Most seeds are removed these ecotypes are difficult to germinate. Means to by birds or other animals (Brinkman 1974g). stimulate or assure field germination are not well Fruits may be harvested over a period of 2 to 3 understood, and until these problems are corrected, months during early fall and winter, but they are more use of skunkbush sumac will be adversely affected. difficult to clean if they have been allowed to dry on Seeds that have been properly pretreated will nor- the shrub. Fruits are collected by hand picking or mally germinate in 30 to 45 days, once germination flailing the stems after leaves have fallen in early begins (Monsen and Christensen 1975). Germination winter. Fruits adhere to the plant and are not easily is usually completed in 30 days under nursery condi- dislodged. This characteristic interferes with seed tions (Brinkman 1974g). This long germination pe- collection and results in high seed costs. Harvested riod can adversely affect survival of wildland fruits are macerated using a Dybvig cleaner and plantings. Soil at many sites does not remain moist flushed with water to remove the pulp. The remaining long enough to ensure germination and seedling material, including the seeds, is then dried and fanned establishment. to remove small debris. Seeds can be cleaned to puri- Different ecotypes of skunkbush sumac occur ties exceeding 90 percent. There are approximately throughout its range. Adapted seed sources must be 20,000 cleaned seeds per lb (44,100 per kg) (Brinkman selected for range seedings or transplanting projects. 1974g; Plummer and others 1968; USDA Forest Ser- Scarified seed should be fall planted to provide over- vice 1948). Recommended laboratory standards for winter wet prechilling; scarified seeds sown in the seed purchases are 40 percent germination and 95 spring must first be wet prechilled. Seed should be percent purity. Cleaned seeds remain viable for up to planted approximately 0.5 inch (1.3 cm) deep or slightly 5 years under open, dry storage. deeper in dry, coarse-textured soils. Skunkbush sumac Seed production varies among sites and ecotypes. may be seeded at the rate of 2 to 4 lb per acre (2.2 to 4.5 Most upland populations produce some seeds each kg per ha), depending on row spacings. It may be year, but abundant crops must be produced to justify broadcast on rough surfaces or in pits. If drill seeded, commercial harvesting. Seeds from many useful it should be planted with other slow-growing shrub ecotypes or collection sites normally are not harvested species in rows separate from faster growing species. or sold because production is quite low. Consequently, Seedlings grow moderately well, but young plants poor seed-producing ecotypes are not collected and are not highly competitive with herbs (Monsen 1987). used in restoration programs. Seeds from more lim- Consequently, this species should not be seeded to- ited, but productive, stands are normally harvested gether with grasses. Young plants can be stunted by and sold for extensive plantings. Thus, the most adapted herbaceous competition and may not recover (Monsen ecotypes are not always used. 1987). Established seedlings are quite hardy and resil- Seed germination may be restricted by the presence ient. Range or wildland seedings normally require 2 to of a hard, impervious seedcoat and embryo dormancy. 5 years to fully establish. Both forms of dormancy vary widely among seedlots. In the bareroot nursery seed should be planted Seedcoat permeability may be increased by a 20- about 0.5 inch (1.3 cm) deep. A seeding density of 25 minute to 2-hour sulphuric acid scarification (Babb viable seeds per linear ft (82 viable seeds per linear m) 1959; Brinkman 1974g; Glazebrook 1941; Heit 1968a). was recommended by Brinkman (1974g). At the Los A wet prechilling for 30 to 120 days is required to

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 609 Chapter 23 Shrubs of Other Families

Lunas Plant Material Center seeds are planted at a structures. The plant is particularly important for density of 182 per linear ft (600 per linear m) of row to recreational sites in semiarid areas in the Inter- yield 23 usable plants per linear ft (75 per linear m) mountain region and the Southwest. (Frazier 1979). Beds should be mulched to prevent Selected ecotypes of skunkbush sumac have proven excessive drying. Plants may be lifted as 1-0 or 2-0 well adapted to unstable disturbances. It has been stock, depending on their growth rate. Field trans- used to stabilize sand dunes (Kozlowski 1972), road- planted seedlings should be at least 8 to 11 inches (20.3 ways, and mine sites (Hassell 1982; Monsen and to 28 cm) tall. Container stock is most easily estab- Christensen 1975). However, direct seedings and trans- lished from germinants and requires a 4- to 6-month plants establish and grow at only moderate rates for cropping time (Landis and Simonich 1984). Ferguson the first few years. Although survival is often high, and Frischknecht (1981) and Stevens (1981) reported plants do not provide extensive ground cover during that large, well-rooted stock is easily transplanted. the establishment period. Once well established, an- Skunkbush sumac is grown by many commercial nurs- nual growth increases, and the plants begin to provide eries, and wildland plantings of bareroot stock have considerable ground cover and soil protection. been highly successful. Skunkbush sumac can be used to stabilize disturbed Doran (1957) reported that cuttings of Rhus root watersheds, including riparian sites. Its growth habit well when taken in July and potted in a sand-peat provides excellent ground cover, and plants are able to media. This provides a means of propagating collec- resist soil deposition. It is particularly useful for res- tions from specific locations that are difficult to rear toration of meadows and other riparian sites where from seed. erosion and gully cutting have drained wetlands and Field cultivation of skunkbush sumac has been suc- lowered water tables. cessful in improving seed production. Producing seed The spreading moundlike growth form of some from agricultural fields or nursery sites could un- ecotypes and the ability of the shrub to form dense doubtedly provide a means of marketing desired thickets by root sprouting cause skunkbush sumac to ecotypes and providing a more stable market. be a valuable cover plant for birds (Adkins 1980). In addition, the fruits of skunkbush sumac are consumed Uses and Management—Skunkbush sumac has by many bird species (Brinkman 1974g), particularly been extensively used in windbreaks, shelterbelts, during winter months, as berries remain on the bush and conservation plantings (Brinkman 1974g; Hassell (Swenson 1957). Consequently, the shrub has been 1982; Swenson 1957). As an ornamental, it grows selectively planted in windbreaks and associated con- rapidly and develops dense foliage (Johnson 1963; servation plantings to provide forage and cover for Olson and Nagle 1965; Steger and Beck 1973). The birds. leaves and berries turn brilliant shades of red and Skunkbush sumac has been most widely planted in orange in the autumn. Mature plants withstand the Intermountain area for big game habitat. It is burial by drifting sand and sprouting from the crown. often planted in mixtures with other more preferred Plants are also able to survive when soil is eroded from forage plants. The shrub is selectively planted on less the upper portions of the root system. These character- fertile sites in big sagebrush, pinyon-juniper, and istics, along with its cover value, low palatability, mountain brush communities where ground cover and ability to resprout after fire, longevity, and ability to big game habitat are required (Plummer and others withstand climatic extremes, qualify sumac as a valu- 1968). It grows slowly, normally requiring 10 to 20 able species for revegetation (Johnson 1963; Olson years to attain mature stature (Monsen 1987), but it is and Nagle 1965; Swenson 1957). one of the most successful shrubs established from Skunkbush sumac requires little maintenance in rehabilitation plantings. Initial establishment and conservation plantings. Plants do not become deca- ratings of young plantings are regarded as only fair, dent with age, but remain healthy and robust. Many but long-term performance ratings are good to excel- other shelterbelt species often decline in vigor or are lent. Swenson (1957) and Adkins (1980) also reported subject to crown damage from wind, insect attack, or good to excellent survival for 20-year-old cover and snowfall. Skunkbush sumac is generally not adversely windbreak plantings of skunkbush sumac in eastern affected by these agents, but persists and becomes a Colorado and eastern Washington. major species in older plantings. Palatability of skunkbush sumac is rated low for Skunkbush sumac has been widely used in recre- livestock and fair to good for big game, depending on ation sites, parkways, campgrounds, and roadway rest the ecotype and other browse species available (Day- areas. It frequently recovers by sprouting following ton 1931; Stanton 1974). It has been planted in mix- disturbances, and can be managed to assure the tures for range and wildlife habitat to provide forage presence of native species in recreational sites. It with- and cover. In Montana and Wyoming it is considered stands traffic and campground activity; consequently, a preferred browse species and receives heavy winter it is used to screen landscape trails, roadways, and

610 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families use (Mueggler and Stewart 1980; Tweit and Houston mountain brush communities at altitudes between 1980) and moderate use in the summer (Sanford 3,590 and 7,710 ft (1,100 and 2,350 m) (Welsh and 1970). Throughout other portions of its range, the others 1987). Harrington (1964) reported that the plant is only lightly grazed (Sanford 1970). Sheep shrub occupies valleys and slopes throughout the often browse the shrub at all seasons. Skunkbush western portion of Colorado at elevations between sumac is a particularly important forage plant in 5,500 and 7,500 ft (1,700 and 2,300 m). midwinter during periods of deep snow cover. It occu- Rocky Mountain smooth sumac often exists in dis- pies windblown slopes that often remain open and junct stands in dry places, usually in open areas available for browsing. Skunkbush sumac recovers without overstory shade. Large patches or clumps well from wildfires, sprouting vigorously from under- often occur alone without other shrubs, but inter- ground rhizomes (Dwyer and Pieper 1967; Wright and mingled in groupings with other plants. It grows on a others 1979). It is an important shrub in areas where variety of soils including sandy as well as heavy- fires can be used to enhance vegetative conditions. textured materials. It is a climax species with various bunchgrasses in the Northwest, although the native Varieties and Ecotypes—“Bighorn” skunkbush grasses have often been replaced by annual cheatgrass sumac, developed by the Los Lunas Plant Materials (Daubenmire 1970; Franklin and Dyrness 1973). This Center, is a tall, upright variety selected from trials species grows along streams and canyon bottoms align- conducted at locations from New Mexico to Montana ing riparian communities, but does not occur in wet and North Dakota (Thornburg 1982). Seed of this areas. It occupies steep slopes and streambanks inter- variety does not exhibit embryo dormancy and re- mixed with numerous upright trees and shrubs, but quires only a 10 to 25 minute acid scarification (Frazier not as an understory in dense patches of vegetation. It 1979). Populations of skunkbush sumac vary widely in is extremely drought tolerant, and is encountered on size, growth habit, leaf characteristics, and range of open south- and west-facing slopes with big sage- adaptation; they provide many opportunities for fu- brush, antelope bitterbrush, and mountain brush spe- ture selection work. cies. Rocky Mountain smooth sumac also forms thickets in the prairie grasslands (Weaver and Fitzpatrick Family Anacardiaceae______1934). Rhus glabra Plant Culture—Rocky Mountain smooth sumac Rocky Mountain smooth sumac normally produces an abundant seed crop. If grown on favorable sites, plants usually produce heavy seed Description—Rocky Mountain smooth sumac is a crops each year. Even under arid situations, plants small, little-branched, thicket-forming shrub (Van produce some viable seeds. Plants flower from June to Dersal 1938). Mature plants usually do not exceed 3 to August, and are not subjected to late spring frosts that 6 ft (0.9 to 1.8 m) in height. Root suckers develop from the parental plant, and new shoots may appear 10 to 13 ft (3.0 to 4.0 m) from the base of the shrub (fig. 4). Branchlets are glabrous and glaucous. The leaves are mostly 4 to 12 inches (10 to 30 cm) long with 7 to 21 leaflets. Leaves are lanceolate, mainly 0.6 to 3 inches (1.5 to 8 cm) long, acuminate, serrate, and range in color from dark green to light green. The inflorescence consists of a panicle about 4 to 8 inches (10 to 20 cm) long, containing many white flowers. The fruit is a globular, red, thinly fleshy drupe. It is about 0.2 inch (4 mm) long and covered with a dense layer of short red hairs (Harrington 1964; Welsh and others 1987). Ecological Relationships and Distribution— Rocky Mountain smooth sumac is widespread in North America (Welsh and others 1987). It occurs from British Columbia to California, Oregon, and Arizona, and eastward to the Atlantic Coast (Munz and Keck 1959). It is common from Nevada and Mexico to New Hampshire and Georgia (Davis 1952; Hitchcock and Cronquist 1973). In Utah it occurs on a variety of sites Figure 4—Rocky Mountain smooth sumac pro- including desert shrub, riparian, pinyon-juniper, and duces distinctive clusters of red drupes.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 611 Chapter 23 Shrubs of Other Families damage seed crops of many shrubs. The flowers are success. Consequently, harsh sites can be successfully small and borne in terminal or axillary clusters planted with transplant materials. Rooted suckers (Brinkman 1974g). Initially, the developing fruits do can be field dug and transplanted with good results if not appear to be very abundant, but as fruits ripen in lifted when dormant. Doran (1957) recommended root fall or early winter, large, dense clusters of red fruits cuttings be taken in December. develop (Lovell 1964; Plummer and others 1968; Uses and Management—Rocky Mountain smooth Radford and others 1964; Van Dersal 1938). The fruits sumac is a highly useful shrub. It is important for are easily collected by handpicking or beating the wildlife habitat, ground cover, and conservation shrub to dislodge them. Fruits are usually collected in plantings. However, its usefulness in large-scale res- fall after the leaves have dropped and the fruits are toration plantings has been limited by low or erratic more visible. Fruits can persist on the bush until late seed germination. winter without loss of seed viability. Consequently, Plantings of Rocky Mountain smooth sumac on wild- seed collection may be delayed until late winter. land sites have been enhanced by seeding the shrub Once the fruits are harvested they should be cleaned alone or with a few other species in well-prepared to remove the fruit coat and dry pulp. The fruit is a seedbeds. Planting in late fall in areas free of competi- small drupe having a single bony nutlet (Brinkman tive herbs has improved stand establishment. Once 1974g). The fruit is normally dry at the time of established, small seedlings and young plants are very harvest, and the flesh is often left on the seed; how- hardy and survive periods of drought. New plantings ever, the dry fruit makes planting more difficult, are not damaged by moderate grazing, insects, or particularly when mixed with other seeds. When disease, and are able to develop normally under most cleaned, the seeds are more easily stored or seeded. circumstances. Seeds are cleaned using a macerator to grind away Seedling survival can be quite erratic. Some small the pulp. Water is used to float and separate the pulp seedlings are quite vigorous and grow at moderate rates, from the seeds (Brinkman 1974g; Swingle 1939). If the but natural thinning usually results in the loss of a fruits are dry, soaking with water is often required number of young plants. Plummer and others (1968) before maceration. High yields of good seed are nor- ranked this shrub as low to fair for the development of mally obtained from most seed collections. Cleaned mature stands from direct seedings. Seedling losses seeds consist of from 45 to 75 percent of the fruit or usually result from erratic seed germination; many material initially collected (Brinkman 1974g). Seed seedlings emerge in late spring when soil water is rapidly numbers range between 23,000 and 48,000 per lb being depleted. Such seedlings usually do not survive. (50,700 and 105,800 per kg) (Johnson and others 1966; Plants form a single main stem during the first and Lovell 1964; McKeever 1938; Plummer and others second growing seasons. During this time an exten- 1968; Swingle 1939). sive root system is developed. Plants that survive the Seed germination may be erratic (Monsen and first growing season are quite persistent, and few Christensen 1975), and inhibited by a hard impervi- plants succumb after that date. Plants reach maturity ous seedcoat (Heit 1967a; Johnson and others 1966) quickly; within 6 years the shrub is sufficiently devel- and a dormant embryo. Pretreatment using sulfuric oped to produce flowers and seed. Spread by under- acid or hot water to increase seedcoat permeability ground rooting usually begins within 10 years after has been employed (Johnson and others 1966), but planting. Thereafter, root suckering may occur each mixed results have occurred due to differences in year, but the pattern is somewhat erratic. Numerous structure of the seedcoat (Brinkman 1974g; Heit 1967a; suckers may appear in some years, but not in others. Lovell 1964). Seeds can be wet prechilled at 40 F ∞ The size and area occupied by a clump of plants tend (4.4 C) for 2 months (Brinkman 1974g) to relieve ∞ to stabilize within 20 to 30 years. Natural spread embryo dormancy. may continue after this time, but maximum spread is Seeds should be planted in late fall or early winter at usually attained within this time span. Natural spread a depth of 0.75 to 1.5 inches (2 to 4 cm). Some seedlots is not always restricted by the presence of other spe- germinate much better than others, particularly if cies. New shoots often emerge in areas occupied by pretreated. Attempting to predetermine planting closed stands of native bunchgrasses, big sagebrush, success from new seedlots is difficult. Nursery seedings and antelope bitterbrush. are often heavily sown to assure establishment of Rocky Mountain smooth sumac is an important satisfactory stands. forage plant for wildlife. The fruits are eaten by a large Success from seeding wildland sites has been ex- number of birds (Miller and others 1948; Stanton tremely erratic due to poor or very erratic germination. 1974). It is moderately browsed by big game in the Excellent quality transplant stock can be grown as spring, fall, and winter months, but is more heavily bareroot or container stock. One-year-old bareroot grazed in the summer (Hill and Harris 1943; Kufeld stock is easily cultured and field planted with good and others 1973; Nemanic 1942; Plummer and others

612 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

1968; Smith 1953; Snyder 1937). When this shrub long as broad, glossy on the upper surface and dull occurs on big game wintering ranges and is available, beneath (fig. 5), with more pronounced spiny tips than it is heavily browsed when access to other species is Oregon grape. The species frequents moist, rich hu- restricted by deep snow. Some populations are more mus soils on the floor of conifer forests, but sometimes heavily grazed each year than others. grows on exposed rocky slopes, brushy hillsides, and in Rocky Mountain smooth sumac is relatively easy to aspen or alder stands. Marchant and Sherlock (1984) establish by transplanting on harsh, unstable sites report that it is rather slow growing during the first (Plummer and others 1968). Consequently, it has been season and slow to establish on wildland sites. The successfully used to plant mine disturbances, road- species integrates with Oregon grape in northeast and ways, and unstable watersheds. Although it estab- north-central Washington, thus the two taxa may be lishes well and spreads to provide soil stability, erect varieties of a single species. open stands usually develop on semiarid sites. Plants Distribution—Shining barberry is distributed from growing on more mesic sites develop dense protective British Columbia and northern Washington to north- ground cover. Under most conditions, this species can eastern Idaho, and south from the eastern edge of the be used to provide soil protection, particularly if un- Cascade Mountains to the coast, as far as the southern derstory herbs are also planted. Willamette Valley. Rocky Mountain smooth sumac is a useful shrub to plant on unstable slopes, streambanks, and roadfills Plant Culture—Shining barberry has not been where mass movement may occur. Plants naturally extensively used in wildland plantings. Transplants spread by roots to furnish excellent site stability. have been grown for small plantings, but direct seedings Unstable soils can be quickly stabilized by transplant- are rare. Plants normally produce some fruits or ber- ing large stock (2-year-old plants) 2 to 3 ft (0.6 to 0.9 m) ries each year, and seeds can be collected and pro- apart. The plants spread rapidly and fully occupy the cessed as described for Oregon grape. Seeds are diffi- area in 1 to 3 years. cult to germinate uniformly, and information is needed Rocky Mountain smooth sumac has been used to to properly culture this species. revegetate mine disturbances, as it will grow in soils Uses and Management—Shining barberry is in- with low fertility. It does not establish or compete very frequently used for low-maintenance landscaping, well if sites are heavily seeded to herbs and fertilized erosion control, and conservation plantings. It is grown at high rates. by some commercial nurseries, but is not commonly This shrub is widely used as an ornamental because used in most wildland restoration projects. of its low growth form, attractive fruits, and fall coloration. It is used to stabilize and beautify steep Varieties and Ecotypes—None. banks, roadways, parks, and recreation sites. It has been cultivated since 1620 (Brinkman 1974g). Rocky Mountain smooth sumac is also useful in windbreak and conservation plantings. It withstands extreme cold temperatures, and can be used on exposed, windblown sites to protect soils and entrap snow. Rocky Mountain smooth sumac survives wildfires, and is an important plant where burning may create erosion and watershed problems. This species can recover quickly from fire, and provides needed cover the first year after burning. Along with Gambel oak, maples, Saskatoon serviceberry, and chokecherry, the species readily resprouts and protects steep watersheds. Varieties and Ecotypes—None.

Family Berberidaceae ______Mahonia aquifolium Shining barberry Description—Mature plants are 0.3 to 6.6 ft (0.1 to Figure 5—Glossy, elongate, spine-tipped leaves 2.0 m) or more tall. Leaflets are more than twice as characterize shining barberry.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 613 Chapter 23 Shrubs of Other Families

Family Berberidaceae ______exchange of water and air. In addition, most seeds require some wet prechilling to germinate. Mahonia fremontii Both container and bareroot nursery stock have Fremont barberry been grown for restoration plantings. Plants have been reared and used in cultivated gardens and nursery Description—Fremont barberry is a large shrub of sites. With some irrigation and culture, plants attain dry slopes and ridges in pinyon-juniper types of the large stature, provide distinct gray-green foliage, and Southwest. It grows 3 to 12 ft (0.9 to 4 m) tall and form an attractive display of red fruits. produces small, compound, spiny leaves and dark blue or red berries. This shrub has yellow wood (fig. 6). Uses and Management—This shrub has been used in some recreation plantings, but has potential to Ecological Relationships and Distribution— restore disturbances in the southwest shrub complex. Fremont barberry is distributed across the Southwest from western Texas through Sonora to Baja California Varieties and Ecotypes—None. and north, and from California through southern Utah and Colorado. It grows on deep sandy soils in Family Berberidaceae ______valley bottoms and draws. It is also encountered amid rocky outcrops and steep canyon slopes. A number of Mahonia repens individual bushes often grow together in small Oregon grape, creeping barberry, patches, although this shrub does exist interspaced mountain holly, creeping mahonia with other woody species. It is found in the warmer drier regions of southern Utah and northern Arizona. Description—Oregon grape is a perennial creep- It grows intermixed with Utah juniper, Stansbury ing shrub with evergreen hollylike leaves (fig. 7). It cliffrose, blackbrush, and Nevada ephedra. It is com- seldom grows taller than 1 ft (30 cm). The shrub mon in draws and washes that may receive infrequent spreads by stolons, rhizomes, and stem layering; indi- runoff. vidual plants sometimes attain a diameter of 4 to 6 ft (1.8 to 3.3 m). Leaves are evergreen, leathery, and Plant Culture—Berries are generally not produced pinnately compound with five to seven (rarely three each year. Special climatic conditions must occur to or nine) leaflets that are less than twice as long as promote flowering and seed production. However, broad. Leaflets are spinulose, glossy to dull on the bushes planted in more mesic situations in central upper surface, and dull to glaucous on the lower Utah have consistently produced abundant crops surface. Perfect, regular flowers develop in fascicled most years. bracteate racemes in the leaf axils. The perianth Seeds are formed in a berry that dries at the time of consists of five alternate whorls of three members maturation. Seeds are large and easily extracted and each; all are yellow. Those in the outer whorl are tiny separated from the dry fruit. A hard seed coat restricts bracts; the next two whorls are the sepals, and the

Figure 6—Fremont barberry produces attractive red Figure 7—Oregon grape, a perennial creeping shrub, fruit and small, leathery evergreen leaves. It grows in occurs on well-developed soils as an understory in a variety of southwestern shrub communities. conifer forests.

614 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families inner whorls are the petals. Petals are bilobed with Seed and dried berries may be fall sown in the field two glands at the base. There are six stamens and a or nursery, or wet prechilled seed may be planted in one-celled ovary. Fruit is a one- to several-seeded, spring. Fall seeding is preferred. Seedlings are vigor- deep blue, glaucous berry about 0.3 inch (8 mm) in ous. Bareroot seedlings may be transplanted after one diameter (Harrington 1964; Welsh and others 1987). or two seasons of growth in nursery beds. Container Oregon grape growth is intermittent in winter and plants can also be established from seed (Landis and more active in spring (Wasser 1982). Flowering occurs Simonich 1984). from March to July. Fruits ripen from September to Seed germination is erratic and may lead to unpre- October and may remain on the plant into winter. Seeds dictable stands, although establishment from plant- are dispersed by birds and mammals (Rudolf 1974). ing stock is usually quite good. Creating depressions for water catchments around planted seedlings, and Ecological Relationships and Distribution— watering immediately following planting may be help- Oregon grape ranges from British Columbia to Alberta ful, particularly on rocky sites. and North Dakota and southward on the east side of Plants may be propagated vegetatively from cut- the Cascade and Sierra Mountains through the Great tings taken from spring through fall and rooted on Basin, Arizona, and New Mexico. It grows with juni- mist benches (Hartmann and others 1990). Best re- pers, pinyon, aspen, mountain brush, lodgepole pine, sults are obtained if basal cuts are made between the spruce-fir, and ponderosa pine at elevations ranging nodes about 1.2 inches (3 cm) above the base of the from near sea level to 10,000 ft (3,000 m) (Harrington current year’s growth (Doran 1957). Rooting may be 1964; Hitchcock and others 1964; Wasser 1982; Welsh hastened by treatment with 5,000 ppm IBA. Layers and others 1987). It is abundant in coniferous forests are occasionally used for vegetative propagation. receiving more than 15 inches (38 cm) of annual Transplants are generally used instead of direct precipitation, and is fairly common on north-facing seeding for landscape and erosion control purposes slopes and other moist sites in plains, foothill to reduce the time until plants reach mature size. shrublands, and low-elevation woodlands. Oregon Nursery-grown stock and wildings are easily trans- grape tolerates full sun and partial to deep shade of planted. Large stock is more difficult to transplant shrubland and coniferous forests. It requires well- due to the development of thick stolons. Commercial drained, more-or-less neutral (pH 5.5 to 7.0) soils, and planting stock is widely available, but the seed or is only weakly tolerant of salinity (Stark 1966). Or- transplants should originate from an area with egon grape is intolerant of poor drainage and high characteristics compatible with those of the plant- water tables. Although it grows in deep, medium- ing site. textured soils, it is also common on shallow, rocky sites (Bailey 1949; Harrington 1964; Plummer and others Uses and Management—Oregon grape has been 1968; Wasser 1982). used in landscape planting as an evergreen ground Plant Culture—Good fruit crops occur erratically. cover. It is quite useful because of its colorful flowers, Fruits are most often collected by hand. Berries of berries, and leaves. Under cultivation Oregon grape more upright populations can be beaten into con- will grow 2 ft (61 cm) tall. The prickly, hollylike leaves tainers (Rudolf 1974). Pulp is removed by macerating are brilliantly red to bronze colored in winter and are the fruit in a Dybvig cleaner. The seeds and pulp are used in Christmas decorations. Its spreading habit dried, lightly chopped, and separated by screening. and ability to grow on fairly dry, exposed, rocky slopes, Seed may be stored dry in air-sealed containers at 34 to make it a good candidate for mined lands, game ranges, highways, and recreation areas. Ecotypes grow- 37 ∞F (1 to 3 ∞C) for 4 to 8 years without significant loss of viability (Heit 1967c). Seeds kept under uncon- ing as an understory with ponderosa pine and other trolled warehouse conditions have remained viable conifers are, however, not well suited to dry, exposed sites. for 13 years (Stevens and Jorgensen 1994). There are The forage value of Oregon grape has been rated as 71,000 seeds per lb (156,600 per kg) at 100 percent nearly worthless for livestock and wildlife in summer purity. Minimum purchasing standards are 95 percent (USDA Forest Service 1937). Smith (1953) ranked it purity and 85 percent germination. last of 33 shrubs tested as summer forage for mule Seed dormancy is complex and variable. A wet deer in northern Utah. However, both livestock and wildlife use the plants when other vegetation is scarce, prechilling period of 1 month at 34 ∞F (1 ∞C), followed by 2 months at 70 F (21 C), and 6.5 months at 34 F particularly during winter (Monsen and Christensen ∞ ∞ ∞ 1975). Kufeld (1973) found Oregon grape valuable for (1 ∞C) relieves dormancy of some seedlots (Rudolph 1974; Wasser 1982). In some cases a 5-day water soak Rocky Mountain elk in fall and winter but of low value may successfully substitute for wet prechilling (USDA in spring and summer when other vegetation was Forest Service 1972). Incubation of excised embryos at available. Mule deer use it to a moderate extent in winter, spring, and fall (Kufeld and others 1973). Patton 68 ∞F (20 ∞C) for 10 to 14 days is recommended as a relatively rapid means of testing germination. and Ertle (1982) reported that in the Southwest it

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 615 Chapter 23 Shrubs of Other Families receives greatest use by elk in winter; mule deer use it rugosa representing the American plants. Thinleaf in the spring and fall. Plants are quite tolerant of alder, also referred to as mountain or river alder, is one browsing. Once frosted in fall, Oregon grape berries of approximately 30 species of this genus that occurs in become more palatable and are used by birds and North America, Europe, and Asia (Schopmeyer 1974a). small mammals (Thornburg 1982). The species is also It is widespread from Alaska and the Yukon south to considered desirable for honey production. Plants con- New Mexico and California, usually growing at eleva- tain several alkaloids, including berberine, which has tions between 5,000 and 10,000 ft (1,500 and 3,000 m) medicinal uses (Suess and Stermitz 1981). The inner in the southern portions of its range (Harrington bark and wood of barberris species are yellow and 1964). were used by Native Americans to prepare a yellow Like most other alders, thinleaf alder occurs in dye. mountainous regions, often growing along streams Seedlings grown on adapted sites may be vigorous, and in wetlands. It is adapted to drier sites and grows but require several years to reach mature size. Control as an understory with various conifers. It spreads of rodents and rabbit populations has been necessary naturally onto disturbances and clearings caused by to establish young plants in shelterbelts in the Mid- wildfires, road construction, and related activities. At west, as Oregon grape is preferred by cottontail rab- lower elevations and on drier sites, thinleaf alder is bits (Swihart and Yahner 1983). Mature plants resprout more restricted to wet areas along streams, seeps, and following burning, and are resistant to browsing. moist mountain slopes (Arno and Hammerly 1977; Hansen and others 1988a; Komarkova 1986). It does Improved Varieties—None. A number of released varieties have been developed for the ornamental trade.

Family Betulaceae ______Alnus incana Thinleaf alder Description—Thinleaf alder is a deciduous, multistemmed shrub or small tree (fig. 8). Plants are usually 3 to 13 ft (0.9 to 4 m) tall (Welsh and others 1987), but may attain heights of 40 ft (12.2 m) (Arno and Hammerly 1977; Patterson and others 1985). The bark is gray to brownish (Patterson and others 1985; Preston 1948); twigs are puberulent and commonly glandular (Welsh and others 1987). Leaves are broadly elliptic or ovate-oblong, 0.4 to 1.2 inches (1 to 3 cm) long with a dull green color and doubly dentate margins (Hitchcock and Cronquist 1973; Patterson and others 1985). Male and female flowers are borne in separate catkins on the same plant. The staminate catkins are clustered at the ends of twigs, each 1 to 4 inches (2.5 to 10 cm) in length (Hitchcock and Cronquist 1973). The pistillate catkins arise from branches of the previous season (Welsh and others 1987) and occur in groups of three to nine on the ends of branches. These eventually develop into small cones that are 3.6 to 5.2 inches (9 to 13 cm) in length (Hitchcock and Cronquist 1973; Patterson and others 1985). The fruit is a small single-seeded nutlet with narrow wings (Haeussler and Coates 1986; Mozingo 1987; Schopmeyer 1974a). Ecological Relationships and Distribution— Welsh and others (1987) recognized thinleaf alder as a portion of a huge circumboreal complex, with Alnus incana spp. rugosa var. occidentalis being the Old Figure 8—Small multistemmed trees of thinleaf alder World portion and Alnus incana spp. rugosa var. border a small stream in eastern Idaho.

616 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families extend into big sagebrush communities at lower eleva- carried considerable distances with strong winds. Seed- tions growing along streams, seeps, and springs. It is lings developing from seeds broadcast on rough seed- most prevalent in the Douglas-fir, ponderosa pine, beds establish well if the surface does not dry rapidly. spruce-fir, and lodgepole pine communities within the Seedlings do not establish well on compacted surfaces Intermountain region. It occupies sites that may be and impervious soils. Thinleaf alder has been broad- seasonally flooded or where water tables remain near cast seeded in a slurry using a hydromulch, but with the soil surface. Although often restricted to areas limited success. Better stands have been achieved by with high water tables, it is also found on well-drained dry seeding. sites. It is often abundant as an understory with Thinleaf alder seeds can be planted with most drill conifers, as it is shade tolerant (Haeussler and Coates seeders. If seeded alone in nursery beds, the seedlot may 1986; Hansen and others 1988b; Kauffman and others require dilution with inert material to aid in regulating 1985; Kovalchik 1987; Padgett and Youngblood 1986; the seeding rate. However, many precision nursery Youngblood and others 1985). seeders can plant pure alder seeds. Seeds should be Thinleaf alder grows on well-developed, highly or- placed about 0.25 inch (0.6 cm) deep on firm, but ganic soils (Kovalchik 1987), but occurs on well-drained, noncompacted, seedbeds. Planting on seedbeds having poorly developed, cobbly gravels and sandy textured considerable surface litter has been quite successful. soils (Hansen and others 1988a; Kovalchik 1987). This Within its range of climatic adaptation, thinleaf shrub is able to invade flooded or burned sites (Crane alder will invade fresh disturbances. It responds as a 1982; Zasada 1986). It responds as an early seral pioneer plant capable of establishing in open areas species (Kauffman and others 1985; Zasada 1986), and (Zasada 1986). It is able to compete well with herba- is very often abundant following disturbances. It is ceous species when seeded on both disturbed and commonly encountered dominating disturbed sites undisturbed sites. Seedlings develop quickly and can including roadways and riparian habitats. establish amid considerable herbaceous competition. New shrub seedlings often invade roadway and mine Plant Culture—Both staminate and pistillate cat- disturbances where seeded herbs have previously es- kins develop during the growing season and persist tablished. On infertile soils, thinleaf alder may grow through most winter months. The catkins begin growth more vigorously than the herbs. early in spring before leaves develop. Flowering occurs Thinleaf alder is not universally adapted to all soil in early March and April, depending on elevation and disturbances and it is particularly sensitive to compe- site location. Fruits ripen in early fall (Schmidt and tition from some species. Although the shrub invades Lotan 1980). Fruits are small winged nutlets borne in areas seeded to grasses, its growth can be suppressed pairs on bracts of small cones or strobiles—when by some herbaceous species, particularly smooth brome. dispersed they are spread by wind or water (Haeussler Planted seedlings that do not become inoculated with and Coates 1986; Mozingo 1987; Schopmeyer 1974a). soil organisms remain small and can be suppressed. Thinleaf alder normally produces abundant seed crops Although these factors are not well understood, most years. Some individual plants bear heavy crops irregularities in plant performance on disturbed sites every year. There are about 675,000 cleaned seeds per can be expected. lb (1,488,000 per kg) (Schopmeyer 1974a). Seed viabil- Thinleaf alder can be easily propagated and planted ity is quite variable. Schopmeyer (1974a) reported as bareroot nursery stock or container material (Platts some collections contain less than 5 percent viable and others 1987). Transplants usually establish quickly seed. However, good quality seed can be obtained by and grow considerably during the year of planting. collecting from healthy bushes early in the season One-year-old transplants develop fibrous root sys- before seeds shatter. Bushes with disfigured cones or tems that greatly enhance survival. Thinleaf alder is infested with disease should be avoided. Also, seeds reported to fix nitrogen (Haeussler and Coates 1986). from plants that are weakened by climatic stress Plants inoculated with nitrogen-fixing organisms begin should not be harvested. Seed samples should be growth soon after planting. Other plants that appar- cleaned and tested for germination before seeding ently lack these organisms grow slowly, but respond rates are determined. Seeds should not be stored in dramatically if inoculated. Adding soil containing natu- open warehouses for more than 2 years because viabil- ral microflora to containerized stock appears benefi- ity decreases rapidly after this time. If possible, air- cial. When plants are established on harsh sites, dried seeds should be stored in sealed containers at 34 fertilizer applications have benefitted initial growth. to 38 F (1 to 3 C) (Schopmeyer 1974a). Seeds are ∞ ∞ Seedings of thinleaf alder on sites supporting a herba- nondormant when collected and do not require wet ceous cover are often successful in conifer forest com- prechilling to germinate (Haeussler and Coates 1986). munities. This shrub has frequently invaded roadways, Thinleaf alder seeds are small, lightweight, and logging disturbances, and mine sites that were first easily dispersed. Broadcast seeding, particularly using seeded to introduced and native herbs (Plummer 1977). aircraft, may create irregular stands, as seeds can be

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 617 Chapter 23 Shrubs of Other Families

The shrub invades seeded areas where established Collections of thinleaf alder do not appear to be as herbs are weakened due to decreases in soil fertility. site specific as many other shrubs. Plantings have Its seedlings apparently are very competitive and been successful over a range of elevations and site vigorous. Once established, few other plants grow as conditions. The species should not be planted on sites rapidly. where it does not normally occur, but it can be used on mine or road disturbances within its natural range of Uses and Management—Thinleaf alder is par- occurrence. ticularly useful in providing cover and forage for wild- Thinleaf alder plants are not usually lost to insect life. Abandoned logging roads and other disturbances damage, climatic stress, or herbivore grazing. The spe- that are seeded to grasses often are invaded by thinleaf cies is long lived and compatible with overstory trees. alder, snowbrush ceanothus, chokecherry, and other It is able to exist in open or shaded areas and can be important browse plants. Game animals seek these used as a low maintenance species in recreational areas to graze the variety of plants. Animals also plantings. Thinleaf alder is an attractive shrub and utilize thinleaf alder in burned and cutover lands; can be used as a screen or specimen plant for horticul- plants quickly recover and produce an abundance of ture plantings. It remains vigorous and green the herbage. Use by big game animals varies among sites. entire summer and may be used to reduce fire spread Light to moderate use has been reported by deer, elk, around recreational sites. and moose (Heale and Ormrod 1982; Kauffman and This species has been overlooked in many wildland others 1985; Knowlton 1960, Kufeld 1973). However, restoration programs. Although recognized as a rapid- Kufeld and others (1973) reported that alder is an developing species capable of producing considerable important browse for mule deer. Beaver and other biomass, it has not been emphasized for watershed or small animals and birds also utilize this species (Arno wildlife plantings. and Hammerly 1977; Rue 1964). Thinleaf alder provides excellent cover for wildlife. It occupies riparian Varieties and Ecotypes—None. zones, wetlands, and brush fields where animals concentrate. The establishment and growth traits of this species cause it to be one of the most useful shrubs for Family Betulaceae ______revegetation of riparian sites in the Intermountain Betula glandulosa region. Bog birch Varying degrees of use by cattle and sheep are reported for thinleaf alder (Dittberner and Olsen 1983; Description—Bog birch is a deciduous, highly vari- Roath and Krueger 1982). Animals selectively graze able, spreading to erect monoecious shrub ranging certain stands more heavily than others. Use is some- from 0.3 to 19.7 ft (0.1 to 6 m) tall. Plants have one to what regulated by access, acceptance of associated many main stems and deep, spreading root systems. plants, and season of grazing. Livestock graze thinleaf Alternative common names are glandular, resin, alder very heavily in many riparian areas. The plant swamp, marsh, and dwarf birch. Young branches are is frequently maintained in a stunted condition, but is densely puberulent and resinous with yellowish able to recover when grazing is well managed. New wartlike, crystalline glands. Bark is reddish-brown to plantings can be destroyed by heavy grazing. gray or purplish and does not readily peel. Lenticels Thinleaf alder is particularly useful as a ground are rarely conspicuous. Leaves are alternate, oval to cover and erosion control plant. It is one of a few shrubs orbicular, elliptic, or sometimes obovate or ovate. They that, as a seedling, is able to compete with seeded are 0.4 to 0.8 inch (1 to 2 cm) in length, but range from herbs, and can be seeded in mixtures. It grows rapidly 0.2 to 1.6 inches (0.5 to 4 cm) long, with finely serrate and provides cover and herbage within a few years. It or crenate-serrate edges (fig. 9). Blades are bright is well adapted to a wide array of soil conditions, green, thick and leathery, glandular on both surfaces, particularly infertile sites. Its ability to fix nitrogen glabrous above, and paler and puberulent beneath. apparently contributes to its adaptability to harsh Staminate catkins are elongate and pendulous in situations. The presence of thinleaf alder benefits the clusters of one to four. They develop in summer and growth of understory herbs, and thus improves ground flower with or before leaf development the following cover and stability on erodible sites. When trans- spring. Staminate flowers are three per cluster, each planted onto open disturbances it grows rapidly and is with three perianth segments and two stamens sub- able to stabilize sites where seeding is difficult. tended by three bracts. Pistillate catkins appear with Thinleaf alder has been used successfully to reveg- the leaves. They are conelike, erect, and solitary with etate road and logging disturbances in the Idaho two or three flowers per bract. Fruits are narrow- Batholith and on a number of mine sites. It has spread winged, single-seeded samaras. Flowering occurs from onto abandoned mine disturbances, including rocky April to September. Samaras ripen from July to Octo- dredge piles and exposed substrata. ber (Brinkman 1974c; Hitchcock and others 1964;

618 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

associated with bog birch include rushes, horsetails, soft-leaved sedge, beaked sedge, bluejoint reedgrass, and tufted hairgrass (Hitchcock and others 1964; Kovalchik 1987; Patterson and others 1985; Welsh and others 1987; Youngblood and others 1985). Bog birch commonly grows in fens, swamps, or bogs having large accumulations of organic matter derived from sphagnum or nonsphagnum plant materials. On flood plains it often grows in soils with textures ranging from silty to fine sandy loams to organic loam textures (Kovalchik 1987). Mineral requirements of bog birch may vary with population. Krajina and others (1982) and Moss and Wellner (1953) reported the species occurs in acidic or humic soils that are low in nutrients, particularly calcium and magnesium. Figure 9—Stems of bog birch have reddish to purplish bark and thick, serrate leaves. They noted the absence of bog birch from swamps and fens with high nutrient contents derived from adjacent uplands. However, Pojar and others (1984) noted the reverse to be true. Plant Culture—Most samaras are wind dispersed Viereck and Little 1972; Welsh and others 1987); they in fall. Smaller quantities are released through the are dispersed by wind, gravity, and sometimes water. winter and early spring as the catkins disintegrate on They may also be blown for some distance across the shrub. Many of these may be consumed by birds. crusted snow. Throughout much of the species range, Samaras are sometimes collected from the surface of reproduction is primarily by seed. In the northern- crusted snow. Fruits are harvested, processed, and most limits of its range, seedlings are rare and plants stored as described for water birch. There are between spread primarily by layering (Hermanutz and others 3 and 5 million cleaned seeds per lb (6,600,000 and 1989; Weis and Hermanutz 1988). 11,000,000 per kg) (Brinkman 1974c). Culture is as described for water birch (Brinkman 1974c; Lotan Ecological Relationships and Distribution— 1981). Kelly (1970) reported that bog birch’s value in Bog birch is widely distributed from the interior of rehabilitation is limited; it does not transplant easily. Alaska across northern Canada to Labrador and In the northern portion of the species range, few viable Greenland. In the West, it is found in mountainous seeds are produced, and bog birch spreads primarily areas from coastal British Columbia to California by stem layering (Hermanutz and others 1989; Weis and Colorado at elevations from 4,000 to 11,000 ft and Hermanutz 1988). (1,200 to 3,400 m) (Dittberner and Olsen 1983; Harrington 1964; Welsh and others 1987). Bog birch Uses and Management—Bog birch is consumed occurs in wet meadows, fens, swamps, bogs (sphag- by many wildlife species. Moose, elk, and mule deer num and nonsphagnum), muskegs, moist-to-wet tun- make moderate to heavy use of the shrub in summer dra, wet meadows, and along low gradient streams and winter (Dorn 1970; Kufeld 1973; Kufeld and and lakes. It is also found on sites where the water others 1973; Zach and others 1982). Pine siskin, chicka- table remains high from runoff in nearby uplands. Bog dees, kinglets, and many other birds use the catkins, birch maintains itself in these moist habitats and buds, and seeds (Brayshaw 1976; Komarkova 1986; appears to be a climax species under such conditions Pojar and others 1984; Stephens 1973). Livestock (Marr 1961; Pojar and others 1984). In alpine areas it make only light to moderate use of bog birch except in may also be found in dry rocky habitats (Brayshaw late summer when the boggy soils dry out (Dayton 1976; Komarkova 1986). Bog birch is highly frost 1931; Hansen and others 1988b; Kovalchik 1987). tolerant and is widely distributed in permafrost areas Energy and protein value of the species are low. (Krajina and others 1982). Its shade tolerance is mod- Wetland areas supporting bog birch burn only infre- erate to high. quently due to the normally high water content of the Bog birch most often occurs on wetland sites in vegetation and soil (Crane 1982; Kovalchik 1987). lodgepole pine, Engelmann spruce, or subalpine fir Such sites frequently act as firebreaks, but sometimes forest types (Hansen and others 1988b; Kovalchik burn during dry summers or in late fall. Shoot systems 1987; Olson and Gerhart 1982; Pierce and Johnson may be destroyed but the plants resprout rapidly, 1986). It is frequently associated with alder, willows, particularly if the organic layer of soil around the base Woods rose, Douglas spiraea, blueberry, mountain huckle- is not removed. Germination of wind-dispersed seeds berry, and redosier dogwood. Herbaceous species from surviving plants, or seeds arriving from offsite,

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 619 Chapter 23 Shrubs of Other Families might be facilitated or enhanced by the presence of two stamens, all subtended by three bracts. Blooming exposed mineral soils. occurs as the leaves expand. Female flowers develop in conelike catkins 1 to 1.5 inches (2.5 to 4 cm) long. Varieties and Ecotypes—None. Water birch flowers in April and May. Fruits turn brownish tan as they ripen in fall. Samaras are shed Family Betulaceae ______from the catkins in fall and winter and are dispersed by wind, gravity, and sometimes water. They may be Betula occidentalis blown for considerable distances across crusted snow. Water birch Weather conditions and foraging by seed-eating birds contribute to overwinter disintegration of the catkins Description—Water birch, also known as spring or (Brinkman 1974c; Lanner 1983). red birch, ranges from shrublike to treelike in growth habit (fig. 10). Plants grow rapidly and are short lived. Ecological Relationships and Distribution— They may attain heights of 36 ft (11 m), with clump There are more than 40 species of Betula in the North diameters of up to 20 ft (6 m), and major trunk Temperate and Arctic Zones. Some taxa hybridize, diameters of 1 ft (0.3 m). New stems develop from the producing intermediate populations in zones of distri- root crown, eventually forming dense, multistemmed bution overlap (Dugle 1966; Hitchcock and Cronquist clumps with ascending branches and open crowns 1973). Water birch is distributed from Alaska to Cali- (Hansen and others 1988a; Youngblood and others fornia and east to Saskatchewan and New Mexico. It 1985). The thin, smooth bark is dark brown to black on forms dense thickets in riparian woodland communi- young trunks; it becomes lustrous coppery to brownish ties and in moist areas of drier habitats (Arno and red on older trunks. Lenticels are light colored and Hammerly 1977; Hansen and others 1988a; Padgett vertical. The alternate, deciduous leaves are petiolate and Youngblood 1986; Welsh 1974). It is common with bright green stipules. They are glabrous to pubes- along streams, on steep slopes, or on alluvial terraces cent, glandular, ovate to suborbicular or slightly obo- from Engelmann spruce, ponderosa pine, and Douglas- vate, and 1 to 2 inches (2.5 to 5 cm) long. The blade apex fir zones to sagebrush communities (Hansen and others is rounded to acute, the base usually rounded, veins 1988a,b; Padgett and Youngblood 1986; Youngblood prominent, and the margins sharply once or twice and others 1985). Communities tend to remain rather serrate. stable in spite of annual flooding (Padgett and others Male and female flowers develop in separate inflo- 1989; Youngblood and others 1985). However, the rescences on a single plant. Staminate catkins are plants are susceptible to windthrow because they formed during the summer or fall and rapidly elongate grow on sites with high water tables and are usually into tassel-like catkins 1 to 4 inches (2.5 to 10 cm) long shallow rooted (Lanner 1983). In eastern Washington, the next spring. Staminate flowers grow in clusters of Oregon, western Idaho, and southern British Colum- three; each consists of three perianth segments and bia, water birch hybridizes with paper birch, producing many localized intermediate forms (Dugle 1966; Hitchcock and others 1973). It is closely related to bog birch (Betula glandulosa), and in western Canada hybridizes with this species (Dugle 1966). Water birch is listed as a dominant species in a number of communities or habitat types of the Inter- mountain region (Hansen and others 1988a,b; Olson and Gerhart 1982; Padgett and others 1989; Youngblood and others 1985). It occurs with many woody riparian tree and shrub species including cottonwoods, maples, aspen, thinleaf alder, willows, alder, and redosier dogwood (Lanner 1983; Olson and Gerhart 1982; Padgett and others 1989). It is also associated with many shrubs that are facultative riparian species such as Woods rose, Nutka rose, chokecherry, Saskatoon serviceberry, currant, and skunkbush sumac (Hansen and others 1988a,b; Kelly 1970; Olson and Gerhart 1982; Padgett and others 1989; Youngblood and others 1985). Figure 10—Water birch forms attractive foliage Water birch is adapted to soils ranging in texture and flowers as it grows along valley bottoms and from silty to sandy to coarse-textured types that contain riparian sites. at least 35 percent rock fragments. These often have

620 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families thin profiles and overlie river cobbles or other rocky system. It is useful in shelterbelts if adequate water is substrates (Hansen and others 1988b; Padgett and available (Cook 1981), and has some potential as an others 1989). Nutritional requirements for magne- ornamental. The hard wood is used locally for firewood sium and calcium are relatively high (Krajina and and fenceposts (Lanner 1983). others 1982). Dense thickets or corridors of water birch provide thermal, hiding, and travel cover for a wide array of wildlife Plant Culture—Seeds should be collected from species (Hansen and others 1988a). Palatability of selected young, accessible, highly productive trees by water birch to big game is usually low. It may be used hand stripping the catkins into bags before they begin when other browse is limited (Hansen and others to disintegrate. Fruits are sometimes shed on snow 1988b; Kufeld 1973; Kufeld and others 1973). Its and may be hand collected. presence in many riparian woodland communities Catkins should be thoroughly dried prior to extrac- contributes to their structural diversity and provides tion. The tiny samaras are separated from the catkins habitats for many bird species (Lanner 1983; by screening them through a round-holed screen and Youngblood and others 1985). Seeds, catkins, buds, fanning to remove the remaining bracts (Brinkman and sap of water birch are used by many birds including 1974c). Samaras should be dried prior to storage at 36 grouse, redpolls, pine siskin, chickadees, kinglets, red- to 38 F (2.2 to 3.3 C). ∞ ∞ napped sapsucker, and broad tailed hummingbirds Seed fill and quality vary greatly among years. Fill (Brinkman 1974c; Gullion 1964; Platts and others 1987). should be estimated prior to harvest by observing Water birch is not highly palatable to livestock, seeds under transmitted light (Patterson and Bruce although it is used to some extent by sheep and goats 1931). Wet prechilling for 4 to 8 weeks releases embryo (Youngblood and others 1985). Dittberner and Olsen dormancy. Dormancy may also be overcome by germi- (1983) rated energy and protein values as fair. Produc- nating seeds in light (Brinkman 1974c; Yelenosky tivity is often reduced by flooding and sedimentation, 1961). and the nearly impenetrable monotypic stands reduce Plantings are generally established from bareroot or livestock access (Hansen and others 1988a). container stock rather than from seed. Emergence of Water birch is usually associated with riparian seedlings from direct seedings on wildlands is erratic. communities that act as firebreaks and burn only Only 15 to 20 percent of the planted seeds of these during late summers or dry years. Shoots will burn in species usually produce usable seedlings. Seeds planted intense fires. Burned plants generally resprout from in nursery beds or containers must be covered as the root crown, which is often protected to some extent lightly as possible (Hartmann and others 1990). Cov- if the soil is wet. New plants may also develop from ering may not be necessary if the soil surface is kept seed delivered by wind or water from areas outside the moist. Newly emerged seedlings are susceptible to burn. Germination is favored by exposure of mineral damping off at low temperatures (Marchant and soils and reduced competition following fires (Crane Sherlock 1984). They benefit from shade for the first 1982; Hansen and others 1988a,b). 2 or 3 months (Brinkman 1974c). Seedlings develop In Utah, water birch is recommended for landscape rapidly, producing dense root systems, and may be plantings in residential areas developed on historic transplanted after one or two growing seasons mule deer winter range. Many traditionally used land- (Brinkman 1974c). Cuttings are difficult to root, but scape plants in these areas have been heavily browsed leafy cuttings will root in summer if planted under and damaged by mule deer. Water birch is normally glass and treated with IBA (Doran 1957; Hartmann browsed only moderately by mule deer and recovers and others 1990). Low-growing forms are easily propa- quickly (Austin and Hash 1988). gated by layering. The root systems of water birch stabilize banks; over- Seed or cuttings used to produce planting stock hanging branches shade the water and supply organic should be harvested from areas near the planting site. matter, which improve fish habitat (Youngblood and Seedlings should be planted early in spring while others 1985). Soils on steep streambanks may be native birch near the planting site is still dormant. susceptible to erosion, especially along trails used by Seedlings planted on adapted sites with good soil livestock and wildlife. Heavy recreational use associ- water establish well and grow rapidly if protected from ated with fishing may also increase sloughing rates browsing and competition. (Hansen and others 1988a). Stands should, therefore, Uses and Management—Water birch provides be maintained for their streambank stabilization value food and cover for many wildlife species (Kufeld 1973; (Hansen and others 1989; Youngblood and others Gullion 1964; Rue 1964; Stark 1966). It is useful for 1985). streambank stabilization in adapted areas. It grows rapidly, producing a dense, fibrous, but shallow root Varieties and Ecotypes—None.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 621 Chapter 23 Shrubs of Other Families

Family Betulaceae ______hybridize widely in areas where their ranges overlap (Hitchcock and others 1961; Marchant and Sherlock Betula papyrifera 1984; Welsh and others 1987). Paper birch Ecological Relationships and Distribution— Description—Paper birch is a treelike species grow- Paper birch is distributed from Alaska east to ing from 60 to 130 ft (18.3 to 39.6 m) tall and 2 to 3 ft Newfoundland and south across the northern tier of (60 to 90 cm) in diameter from a deep root system States including Washington, northern Idaho, and (fig. 11). Alternative names are canoe, white, and Montana. It occurs in riparian habitats, dry bogs, silvery birch. Trees are distinguished by their grayish swamps, and poorly drained acidic soils with pH as low white to coppery bark with long brown horizontal as four (Marchant and Sherlock 1984). It also grows on lines. The outer bark often peels in long strips; the open slopes, rock slides, forest openings, and distur- inner bark is orange. The leaves are ovate to acumi- bances. It occupies soils with silty loam, sandy, and nate and doubly serrate, usually with tufts of hairs in gravely textures. It is an aggressive pioneer species on the vein axils on the lower surface. Leaves are 2 to 5 logged or otherwise disturbed areas, and rapidly in- inches (5 to 13 cm) long and 1 to 2 inches (2.5 to 6 cm) vades areas where mineral soils are exposed. wide. Male flowers develop in early spring on long, Plant Culture—Culture and management are yellowish, pendulous catkins. Female cones are 1 to 2 largely as described for water birch (Brinkman 1974c). inches (2.5 to 6 cm) long, hanging from slender stalks. Cones are more accessible, abundant, and easily har- Fruits are translucent samaras with the wings slightly vested from young trees. Minimum seed-bearing age broader than the nutlet. Paper birch and water birch is about 15 years (Brinkman 1974c). Catkins may open during dry, cold periods from late fall well into winter; when large numbers of samaras are deposited on the surface of crusted snow. Seeds are cleaned using, a number seven sieve to separate fruits from chaff (Marchant and Sherlock 1984). There are 610,000 to 4,120,000 cleaned seeds per lb (1,344,800 to 9,083,000 per kg); the average is 1,380,000 seeds per lb (3,042,000 per kg). Longevity of seeds dried to a water content of 1 to 5 percent and stored at room temperature is about 1.5 to 2 years. Viability tends to decrease rapidly at greater water contents, even if seeds are kept in cold storage (Brinkman 1974c). Seeds require a wet prechilling treatment of at least 30 days or fall sowing to produce nursery stock. Sur- face-sown seeds exposed to light will germinate without cold wet prechilling. Seedlings are subject to damping off if crowded or if temperatures are cool. They are resistant to frost but not shading (Krajina and others 1982). Propagation from cuttings is not recommended (Marchant and Sherlock 1984). Uses and Management—Seedlings have been used for revegetating riparian sites and forest disturbances, improving wildlife cover, landscaping recreational sites, and in ornamental plantings. Plants have established and survived well, but have not been extensively used in wildland plantings. Young plants grow quite rapidly and compete well with understory competition. Paper birch resprouts following fire and is capable of withstanding both periodic flooding and drought (Marchant and Sherlock 1984). It is browsed to some extent by wildlife, horses, and cattle. Fruits are eaten by many birds. Figure 11—White-barked, multiple-stemmed paper Varieties and Ecotypes—None. birch is often found growing in a forest openings.

622 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Family Caprifoliaceae ______Uses and Management—The orange yellow to orange red flowers and foliage are very attractive and Lonicera ciliosa, Orange honeysuckle could be used in roadside, summer home, and recre- Lonicera involucrata, Bearberry honeysuckle ational landscaping. In northern Idaho this species Lonicera utahensis, Utah honeysuckle has survived serious air pollution created by smeltering operations, while a number of other native shrubs Introduction—There are about 180 honeysuckle have succumbed (Monsen 1975). species distributed across the Northern Hemisphere, Varieties and Ecotypes—None. Africa, Java, and the Philippines (Brinkman 1974f), although only about 20 species occur in North America (Hitchcock and others 1959). All are widely planted for Family Caprifoliaceae ______their fragrant flowers and ornamental fruits. Some species provide food and cover for wildlife and are Lonicera involucrata valuable for erosion control and shelterbelt plantings Bearberry honeysuckle (Adkins 1980; Brinkman 1974f; Plummer 1977; Plummer and others 1968; USDA Forest Service Description—Bearberry honeysuckle is a native 1985a). shrub that is found throughout the Intermountain Different species of honeysuckle possess important West from Alaska to Quebec and south to California, traits that are useful for range and wildland enhance- New Mexico, and Mexico (Viereck and Little 1972; ment. Some of the more important characteristics Welsh and others 1987). It has yellow to purplish include consistent and abundant fruit production, low tinged flowers (fig. 12) and black fruit. There are about stature, cover for birds and small mammals, excellent 326,000 seeds per lb (718,700 per kg) at 100 percent establishment traits from direct seeding and trans- purity. This shrub grows to 2 to 6 ft (0.6 to 1.8 m) tall. planting, rapid growth, and excellent windbreak at- It occurs on fertile soils along streambanks and in tributes and conservation features. riparian sites in the aspen, Douglas-fir, and spruce-fir communities. Plant Culture—All three species described in this Uses and Management—Bearberry honeysuckle chapter produce small berries and seeds. Berries are has been planted with some success in conservation usually dislodged by stripping or beating the stems and wildlife projects. It has not done well when planted when fruits are mature. Seeds are extracted from the on disturbed sites. This species is used by large and berries by macerating the pulp and flushing with small game and livestock during spring and summer water. Once seeds are cleaned, they are easily seeded with conventional seeders. Seedlings develop quite successfully, but most plantings have been completed using bareroot or container transplants.

Family Caprifoliaceae Lonicera ciliosa Orange honeysuckle Description—Orange honeysuckle is a native trailing or twining vine. Stems are branched and may become 20 ft (6.1 m) long (Hitchcock and others 1959). Although plants normally grow in wet sites in aspen and conifer forests, they are also encountered in open areas on rocky and shallow soils. This vine naturally occurs in southwestern Canada, from Oregon to Cali- fornia, and east to Idaho and Montana (Hitchcock and others 1959). Individual plants can spread over large areas on fertile or disturbed soils. They root at the junction of each node if the stem remains in contact with a wet soil surface (Monsen 1975). This species is Figure 12—Bearberry honeysuckle, widespread in easily transplanted from stem cuttings. Western North America, is used in conservation plantings because of its attractive flowers.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 623 Chapter 23 Shrubs of Other Families months. It has potential for restoring disturbances Family Caprifoliaceae ______within its natural range, but its seeds are not easily harvested, thus seed availability is limited. Sambucus cerulea Varieties and Ecotypes—None. Blue elderberry Description—Blue elderberry (fig. 14) is a large Family Caprifoliaceae ______spreading shrub or small tree growing to 10 ft (3.0 m) tall. It has distinctive opposite branches and leaves, Lonicera utahensis pithy young stems, and older grayish stems with Utah honeysuckle irregularly furrowed and ridged bark. Flowers are white and borne in large flat-topped clusters (com- Description and Distribution—Utah honey- pound cymes). Fruits are black to blue (Hitchcock and suckle is a native shrub that grows 3 to 6 ft (0.9 to 1.8 m) others 1959; Welsh and others 1987). The root system tall (Welsh and others 1987) (fig. 13). It has pale is generally fibrous and spreading, but a thick taproot yellow to yellowish-white flowers and red fruit. It may be present (Haeussler and Coates 1986; Welsh occurs from British Columbia and Alberta, south to and others 1987). California and Wyoming. It is fairly common in the Intermountain West within mountain brush, ponde- Ecological Relationships and Distribution— rosa pine, aspen, and spruce-fir communities. Blue elderberry is most common on moist, well-drained sunny sites, but it is also somewhat shade tolerant Uses and Management—This shrub has good fire (Dayton 1931; Plummer and others 1968; Van Dersal tolerance, and is used extensively by livestock and 1938) and occurs at elevations between 4,250 and game during the summer and fall seasons. Utah honey- 8,200 ft (1,300 and 2,500 m) (USDA Forest Service suckle does well under heavy grazing and survives 1937). It ranges from British Columbia and western heavy trampling. It has excellent ornamental charac- Alaska south to California, Arizona, New Mexico, teristics including showy flowers and fruit. The plant Texas, and northern Mexico (Dayton 1931; Harrington establishes well from transplanting and can withstand 1964; Hitchcock and others 1959; Little 1979; Powell considerable traffic and other abuses. It should be 1988). It may be found on a wide range of soil types, considered for use in restoration projects, recreational growing from the sagebrush grass zone to well above areas, summer homes, and administrative sites. the mountain brush and ponderosa pine communities. Varieties and Ecotypes—None. It also grows in openings in aspen and spruce-fir communities (Welsh and others 1987). Blue elderberry is found in canyon bottoms, along streams, and on sites that are wet during the spring. In riparian areas, blue elderberry can occur in dense stands; in other areas individual plants are generally widely spaced. Plants are often abundant in burned areas.

Figure 13—Distinctive flowers and leaves of Utah honeysuckle, an understory shrub that grows with mountain brush, ponderosa pine, aspen, and Figure 14—Blue elderberry growing at the base of conifer forests. a rocky slope in eastern Idaho.

624 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Plant Culture—Blue elderberry generally produces jellies, candy, pies, and sauces (Mozingo 1987; Powell a good seed crop every year. Berrylike fruits are 1988). produced in large clusters. Each fruit contains three to Blue elderberry is grazed by livestock and wildlife five seeds (Hitchcock and others 1959). Fruit clusters species throughout the year (Dittberner and Olsen are easily collected by cutting them from the stem. 1983; Kufeld 1973; Kufeld and others 1973; Martin Seeds are extracted from the fruit by using a Dybvig and others 1951; Robinette 1972). However, use varies macerator. The resulting pulp and seeds are dried, by season (Kufeld 1973; Sampson and Jesperson 1963; then lightly chopped and separated using an air Smith and Hubbard 1954; Steele and Geier-Hayes screen fanning machine. Unfilled seeds can be 1987). Spring and summer use are light. Many mature removed by flotation. At 100 percent purity there are plants grow so tall that by late summer much of the about 217,000 seeds per lb (478,500 per kg). Accept- foliage is out of reach of grazing animals. Palatability able purity is 95 percent and germination 50 percent. and use increase dramatically after the fruit ripens Good viability is retained for up to 16 years (Haeussler and the foliage is frosted (Dayton 1931). Plants are and Coats 1986). heavily used by cattle, deer, and elk in fall; they are Germination of blue elderberry is inhibited by a hard especially palatable to sheep. Use decreases in the seedcoat and embryo dormancy (Plummer and others winter after leaf fall, but plants provide important 1968). Without artificial treatment, germination of herbage. The berries and dried fruit, which often planted seeds may occur sporadically over one to three remain on the shrubs through the winter, are con- growing seasons. Scarification in acid or a 2-month sumed and dispersed by birds and many other ani- warm pretreatment followed by a wet prechilling of mals. Buds are used by grouse, rodents, and big game up to 5 months improve germination (Brinkman animals in winter. The shrub provides cover for big 1974j). Fall seeding is recommended to permit over- game and nongame species, and perching and nesting winter wet prechilling (Plummer and others 1968). sites for birds (Brown and others 1977; Dittberner and Seeds in the soil seedbank often germinate readily Olsen 1983; Gray and Greaves 1984). Blue elderberry following fire (Heit 1967a). is persistent and recovers well from moderate grazing Elderberries can be established by direct seeding in and trampling (Monsen 1984; Plummer and others favorable spots (Plummer and others 1968). Seed may 1968; Van Dyne and Payne 1964). Continual, heavy be hand planted in pits on burns, chained areas, or grazing can be detrimental. Seeded areas should not disturbed riparian sites. Elderberry seed may also be be grazed for two or three growing seasons following planted alone or with other shrubs using Hanson or seeding. Buds and branches may be defoliated by thimble seeders. Elderberries should be planted sepa- grasshoppers during mid or late summer during years rately from grasses to protect the shrub seedlings from when population densities are high. Plants generally competition. Seeds should be covered 0.25 inch (6 mm) recover the following year. deep on a firm seedbed. Blue elderberry exhibits good fire tolerance (Aro Blue elderberry is easily propagated, planted, and 1971) and is able to resprout from the root following established using container or bareroot stock (Plummer fire (Little 1979; Preston 1948; Steen 1965; Van Dersal and others 1968). Plants grow quickly in nursery beds, 1938). Germination of seed buried in the soil is enhanced and often develop a thick taproot and spreading root by fire (Heit 1967a; Morgan and Neuenschwander system. Root pruning can eliminate or reduce this 1988). problem. Blue elderberry can also be established from Varieties and Ecotypes—None. There are many cuttings (Plummer and others 1968). ecotypes within the species. Drought-tolerant, shrubby Blue elderberry shrubs begin growth slowly in early populations associated with sagebrush types do well spring, although rapid vegetative development occurs with only 12 inches (30 cm) of annual precipitation. in June or July. The number of stems and buds, as well Tall, almost treelike populations also occur. Some as total productivity, fluctuate greatly from year to populations are consistently good seed producers, while year. Older branches die back during the winter. New others produce erratic seed crops. growth develops from large vegetative buds on the rhizomes. Uses and Management—Blue elderberry has been Family Caprifoliaceae ______used successfully in soil stabilization (Plummer and Sambucus racemosa others 1968), riparian plantings (Carson and Edgerton Red elderberry 1989; Goldner 1984), streambank stabilization planting (Lines and others 1979), and for revegetation of Description—Red elderberry is a native circum- mined areas (Ferguson and Frischknecht 1985; boreal shrub (Hitchcock and others 1959). Two natural Hungerford 1984; Monsen 1984). Fruits of blue varieties occur in the Western United States: S. r. var. elderberry are gathered and used for making wine, microbotrys has red fruits and is widely distributed,

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 625 Chapter 23 Shrubs of Other Families ranging from Wyoming and northern California south Plant Culture—Red elderberry flowers in mid- to Mexico. S. r. var. melanocarpa produces black summer. Fruits ripen in mid to late September. Good fruits, and occurs primarily in the Rocky Mountains seed crops are produced most years. Fruit clusters are from Canada south to the Pacific Northwest and harvested by cutting them from the stems. Seeds are Wyoming (Welch and others 1987). extracted with a Dybvig macerator. Seeds and pulp Red elderberry has distinctive opposite branches are dried, lightly chopped, and processed through an and leaves. Leaves are pinnately compound, leaflets air screen fanning machine to separate seeds and are five to seven serrate, pointed, 1.2 to 5.9 inches (3 to debris. At 100 percent purity there are about 286,000 15 cm) long, and 0.4 to 2.4 inches (1 to 6 cm) broad. seeds per lb (630,500 per kg). Acceptable seed purity is Plants are fairly short, usually about 3 to 6 ft (0.9 to 95 percent and germination is 50 percent. Success 1.8 m) in height. Stems have warty bark with brown from direct seeding can be erratic (Plummer and pith; they have a rank odor if bruised and crushed others 1968). Germination requires wet prechilling; (Kelly 1970; Viereck and Little 1972). The numerous therefore, fall seeding is recommended. Seed should white to cream-colored flowers are borne in terminal be covered lightly and planted in areas where compe- pyramidal clusters (cymes) less than 3 inches (8 cm) tition will be light for a few years. Red elderberry can wide. The fruit is a red drupe that ripens in September. be propagated by cuttings, either hardwood cuttings Each drupe contains two to four seeds (Hitchcock and started in winter or softwood cuttings taken during others 1959; USDA Forest Servcie 1985; Welsh and spring and summer (Doran 1957; Ritter and McKee others 1987). 1964). Most natural reproduction occurs vegetatively from Ecological Relationships and Distributions— buds on stems, rhizomes, and root crowns following Red elderberry occurs in subalpine, spruce-fir, lodge- fire or mechanical damage (Conrad and McDonough pole pine, and aspen communities (fig. 15). It is dis- 1972; Van Dersal 1938). Some reproduction also oc- tributed across North America from Alaska to curs from seed. Seeds are dispersed by birds and Newfoundland and south to New Mexico and Georgia rodents and can remain dormant in the soil for long (Great Basin Flora Association 1986; Viereck and periods. Seeds scarified by fire often germinate in Little 1972; Worley and Nixon 1974). It prefers full to large numbers during the first season following burn- intermediate sunlight, and is not found in full shade ing (Heit 1967a; Hungerford 1984; Kramer 1984). (Haeussler and Coats 1986). Greatest areas of abundance are in openings where winter snow accumulates Uses and Management—Red elderberry is used and remains late in the spring, riparian areas, mead- seasonally by sheep, cattle, deer, bear, porcupine, ows, parks, and other wet areas. Red elderberry grows and elk (Conrad and McDonough 1972; Dayton 1931; in association with tall and intermediate forbs and Kufeld 1973; Kufeld and others 1973; Martin and grass communities. It occurs singly and in patches of others 1951; Ritter and McKee 1964; Smith 1953; various sizes. Steele and Geier-Hayes 1987; Zager 1980). It is one of the most palatable browse species for elk in Idaho and Montana (Gaffney 1941; Young and Robinette 1939; Zager 1980). Palatability is excellent for sheep following frost. All top growth is consumed or dies to ground level each fall. Fruits mature in late August and Septem- ber. They are consumed by birds, rodents, livestock, and big game. Seed is cached and used by rodents. During winter porcupines and mice feed on the buds and bark (Conrad and McDonough 1972). Red elderberry provides valuable nesting and perching habitat and food in the form of berries and buds for a large number of birds (Denslow 1987; Gullion 1964; Martin and others 1951; Van Dersal 1938). Red elderberry is used to stabilize disturbed and erosion-prone areas, especially on aspen and upper spruce-fir sites, riparian areas, seeps, and other moist sites (Platts and others 1987; Worley and Nixon 1974). Bareroot stock and container-grown stock have been used successfully for planting disturbed riparian ar- Figure 15—Red elderberry frequently occurs in for- eas in aspen and ponderosa pine forests (Platt and ested openings and subalpine communities. others 1987). Red elderberry is used as a colorful

626 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families ornamental (Kelly 1970; Ritter and McKee 1964), and four locular. Generally two locules produce a single should be considered when planting recreational sites, mature seed each. The white pulpy fruits are subglobose summer homes, and ski areas. or ellipsoid and 0.3 and 0.6 inch (8 to 15 mm) in diameter. Seeds are elliptical, flattened, creamy white, Varieties and Ecotypes—None. There is, how- smooth, and 0.16 to 0.2 inch (4 to 5 mm) in length with ever, considerable variation within the species. fleshy endosperm and a tiny, poorly developed embryo (Evans 1974; Hitchcock and others 1959; Hopkins and Family Caprifoliaceae ______Rawlings 1985; Kovalchik and others 1988; Marchant and Sherlock 1984; Viereck and Little 1972; Welsh Symphoricarpos albus and others 1987). Common or white snowberry Ecological Relationships and Distribution— Description—Common snowberry is an erect, multi- Common snowberry occur across the United States branched shrub usually 3.3 to 6.6 ft (1 to 2 m) tall, but and Canada except in the far north of Alaska and ranging from 1.6 to 9.8 ft (0.5 to 3 m) in height (fig. 16). Canada and the Southern States. It naturally occurs Plants are fibrous rooted with the densely branched associated with conifer species, in shade and in open- root system concentrated near the soil surface. The ings with full sunlight. species is slightly rhizomatous with sprouts from the Plant Culture—Common snowberry seeds are white thick, spreading rhizomes forming dense thickets. and average 76,000 per lb (167,500/kg). One hundred lb The slender upright stems are hollow, smooth, and (45 kg) of fresh fruit will yield 3 to 5 lb (1.4 to 2.3 kg) of brownish. Twigs are glabrous and obscurely puberu- cleaned seed. Good seed crops are produced in 4 out of lent. Leaves are opposite, thin, pale green, elliptic to 5 years. Flowering occurs from June through July. elliptic ovate, and entire or with a few coarse, irregular Fruits remain on the plant into winter or until eaten. teeth. They are 0.6 to 2 inches (1.5 to 5 cm) long and 0.4 Fruits are processed through a Dybvig macerator, dried, to 1.4 inches (1 to 3.5 cm) wide with petioles 0.1 to 0.2 and run across an air screen separator. Unfilled seeds inch (2 to 4 mm) long. Leaves on sterile shoots may be are removed by flotation. Germination is delayed by somewhat larger. Racemes are terminal or sometimes dormancy, which can be overcome with wet prechilling in the upper leaf axils. They are dense, subsessile and or scarification with sulfuric acid. Germination will few flowered. The perfect flowers are small, pinkish, occur over a number of years. Young plants grow slowly; and bell shaped. The five-parted corolla is 0.2 to 0.3 however, they can mature and flower in 3 or 4 years. inch (5 to 7 mm) long and nearly as wide. The interior is densely hairy. Corolla lobes range in length from Uses and Management—Common snowberry is one-half to nearly equaling the tube length. Anthers an important forage plant in the West, especially for alternate with the petals and nearly equal the fila- sheep and deer, which make considerable use of it year ments in length. There is a single style. The ovary is around. Elk strip the leaves in the fall. Plants can be lost as a result of excessive use. This species has been used extensively as an ornamental. It can be directly seeded or transplanted in restoration projects. It does not do well on disturbed sites such as mines and road fills and cuts. Varieties and Ecotypes—There are a number of ornamental varieties and selections, but none are available for use on rangelands and disturbed sites.

Family Caprifoliaceae ______Symphoricarpos longiflorus Desert snowberry Description—Desert snowberry is a native, long- lived, deciduous shrub. Plants are low growing, erect to arching, and range in height from 1.6 to 5 ft (0.5 to 1.5 m). The ability of the species to spread by rhizomes Figure 16—Open, branching, scattered, pale-green is poorly known. Plants exhibit wide variation in leaf leaves, and shedding bark are distinctive characteris- and twig pubescence. Young stems are persistent, tics of common snowberry. glabrous to pubescent, and tend to spread at right

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 627 Chapter 23 Shrubs of Other Families angles, giving the plant a thorny appearance. The firm Family Caprifoliaceae ______leaves are simple and opposite, 0.2 to 0.6 inch (6 to 15 mm) long and 0.1 to 0.3 inch (2 to 7 mm) wide, oval to Symphoricarpos occidentalis lanceolate or oblanceolate, and entire or with one or Western snowberry, wolfberry two teeth. The fragrant flowers are solitary, paired in the leaf axils, or in small, few-flowered, terminal Description—Western snowberry is an erect shrub, racemes. The calyx and corolla are five lobed and branching weakly from a woody base and short main regular. The slender corolla tubes are 0.4 to 0.7 inch branches. Plants are 1 to 3.3 ft (0.3 to 1 m) tall, (1.0 to 1.8 cm) long, salverform, glabrous, and pale to spreading freely from rhizomes, and often forming deep pink. Lobes are much shorter than the tube and dense thickets that exclude other vegetation. Roots rather abruptly spreading. Anthers are sessile, alter- are densely branched, shallow, and intermixed with nating with the corolla lobes. The single style is stiff stout creeping rhizomes. Young twigs are puberulent and hairy above the center. The ovary is four loculed; to glabrous. Petioles are 0.1 to 0.4 inch (3 to 10 mm) the two functional locules each contain a single seed. long. The opposite leaves (fig. 17) are thick, elliptic to The fruit is a white, waxy, two-seeded berry about 0.3 ovate, entire or with a few, coarse, blunt, irregular to 0.4 inch (8 to 10 mm) in diameter. Seeds are 0.2 inch teeth. Most are 0.8 to 3.1 inches (2 to 8 cm) long and 0.4 (5 mm) long and acute at the base (Cronquist and to 2 inches (1 to 5 cm) wide with somewhat revolute others 1984; Davis 1952; Evans 1974; Welsh and margins. Leaves on sterile shoots are often larger, others 1987). frequently more lobed, glabrous above, and usually hirsute puberulent beneath, at least along the main Ecological Relationships and Distribution— veins. Few to many-flowered spikelike racemes are Desert snowberry occurs on deep, well-drained soils in produced at the ends of the twigs and in the upper leaf desert-shrub, pinyon-juniper, sagebrush, mountain axils. The five-merous corollas are often wider than bush and ponderosa pine communities from Oregon to long, pink to rose colored, and densely hairy within. Colorado, and south from southern California to Texas. Lobes are arcuate spreading. Anthers alternate with It is the most drought tolerant of the snowberries. It the petals and are shorter than the filaments. The grows intermixed with other species, but not in closed style may be hairy near the middle, but is occasionally stands. glabrous. The ovary is four loculed; two locules usually Plant Culture—Desert snowberry produces white, produce one mature seed each. Fruits are subglobose two-seeded berries. There are an average of 70,000 and greenish white, turning black after frost. The two seeds per lb (154,000/kg). One hundred lb (45 kg) of seeds are white, somewhat ellipsoid, and flat on one fresh fruit will yield 5 to 7 lb (2.3 to 3.2 kg) of cleaned side with a poorly developed embryo (Cronquist and seed. Good seed crops are produced in about 2 out of 5 others 1984; Davis 1952; Hansen and others 1988b; years, depending on spring and summer storms. Flow- Hitchcock and others 1959; Wasser 1982; Welsh and ering occurs in June to mid July. Berries are harvested others 1987). by hand stripping or beating. Fruits are extracted by maceration through a Dybvig cleaner. Unfilled seeds are removed by flotation. Germination can be erratic. Dormancy can be overcome with wet prechilling and scarification with sulfuric acid. Fall seeding will permit prechilling to occur. Young plants can mature and produce seed in 3 to 4 years and reach full stature in 5 to 6 years. Uses and Management—Desert snowberry can be an important browse plant for sheep and deer, and to a lesser extent for cattle. It is fairly resistant to grazing. On many overgrazed sagebrush ranges in which the understory species have been eliminated, desert snowberry and sagebrush will remain. Exces- sive use will result in death of plants. This species can be seeded or transplanted. It does not do well on severely disturbed sites. It does best on soils with well-developed, undisturbed horizons. Figure 17—Western snowberry with ovate leaves and small white flowers is widespread in aspen and Varieties and Ecotypes—None. conifer forests where it often occurs in dense thickets.

628 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Ecological Relationships and Distribution— than wide, elongate campanulate to tubular funnel- Western snowberry is widespread from British Co- form. The corolla lobes are one-fourth to one-half as lumbia to Manitoba, south to Washington, Idaho, and long as the tubular portion of the corolla. The interior Colorado. In Utah it is most often associated with of the tube is sometimes hairy below the level of riparian communities, mainly with cottonwood and filament insertion, but may be glabrous. The fila- willow species (Welsh and others 1987). It does espe- ments are equal to or shorter than the anthers. The cially well on sites with high water tables. It is also ovary is four-locular. Two locules normally contain found in the mountain brush, open conifer sites, and several abortive ovules each; the remaining two loc- upper pinyon-Rocky Mountain juniper communities, ules each contain one normal pendulous ovule. Fruits especially on east- and north-facing slopes. It com- are waxy, white ellipsoid, and about 0.3 to 0.4 inch petes well with understory herbivore species. (7 to 10 mm) long. The hard nutlets or seeds are 0.2 to 0.3 inch (4 to 6.5 mm) long (Cronquist and others 1984; Plant Culture—Fruits of western snowberry are Hitchcock and others 1959; Welsh and others 1987). collected by hand stripping. Seeds are extracted by maceration through a Dybvig cleaner. There are about Ecological Relationships and Distribution— 74,000 seeds per lb (163,000 seed/kg). One hundred lb Mountain snowberry is common in midelevation sage- (45 kg) of fresh fruits will normally yield 8 to 10 lb (3.6 brush, ponderosa pine, Gambel oak, aspen, Douglas- to 4.5 kg) of seed. Good seed crops can be expected 3 out fir, lodgepole pine, and spruce fir communities in the of 5 years. Flowering occurs from the first of June Intermountain region. It can be found growing from through late July. Fruits ripen from mid-September to British Columbia to Montana, and south to California, mid-October. Considerable seed dormancy can exist. Arizona, and New Mexico (Welsh and others 1987). This can be overcome with moist prechilling and Plants may be locally abundant, and in some cases scarification with sulfuric acid. Fall seeding will help form nearly pure stands. Mountain snowberry will to overcome dormancy. Seedling emergence, however, dominate aspen stands. This shrub does well in full can be erratic over a few years. Plants can be estab- sunlight and shade. lished from nursery-grown container and bareroot Plant Culture—Mountain snowberry seeds are stock. Wilding stock can be obtained early in spring collected by hand stripping the two-seeded fruits from while the ground is still moist and before plants break the stems. Good seed crops are produced 3 out of 5 dormancy. years. Seeds are cleaned in a Dybvig macerator. Un- Uses and Management—Western snowberry can filled seeds are removed by floatation. Germination is be used extensively by sheep and deer, and a little less delayed by the impermeable endocarp and an imma- by cattle. Elk prefer it in the fall after it has been ture embryo. Pretreatments include a 3- to 4-month frozen. It can receive heavy browsing and trampling in wet prechilling retreatment. Scarification with sulfu- riparian areas. Excessive use can result in plant death. ric acid is also used to break the dormancy imposed by This species has been used fairly extensively in resto- the endocarp (Evans 1974). The embryo develops to ration projects, especially as transplants. Varieties and Ecotypes—None.

Family Caprifoliaceae ______Symphoricarpos oreophilus Mountain snowberry Description—Mountain snowberry is a spreading to erect branching shrub 4.9 to 6.6 ft (1.5 to 2 m) tall (fig. 18). Leaves and twigs are densely puberulent to glabrous and sometimes glaucous. The thin opposite leaves are elliptic to ovate, entire, or with a few teeth or lobes. They range from 0.4 to 2 inches (1 to 5 cm) in length and 0.1 to 1 inch (0.3 to 2.5 cm) in width. Leaves on sterile shoots may be larger (Mozingo 1987). The cream-colored to pinkish flowers have an unpleasant odor. They are solitary or paired on short, drooping pedicels in the upper leaf axils, or borne on short, few- flowered terminal racemes. The corolla and calyx are Figure 18—Mountain snowberry is commonly five- or occasionally four-lobed. The corolla is longer encountered in open parks and conifer forests.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 629 Chapter 23 Shrubs of Other Families maturity during an extended period of wet and usually acuminate, 1 to 4 inches (2.5 to 10 cm) long prechilling. There are about 75,000 seeds per lb and about two-thirds as wide. They are distinctively (165,000/kg). One hundred lb (45 kg) of fresh fruits pinnately veined and sometimes wrinkled, sparsely will yield 6 to 10 lb (2.7 to 4.5 kg) of seed. Flowering strigillose and greenish on the upper surface, and occurs through June and July. Fruits mature from paler with pubescence along the veins and sometimes mid-September through October. covering the lower surface. Flowers are clustered in Successful establishment may be achieved by fall flat-topped, ebracteate cymes. Penduncles and planting. Seeds may be hand planted or drilled. Seeds branches are strigose to conspicuously spreading pu- should not be directly planted with grasses because bescent. Flowers consist of four very small sepals, four snowberry seedlings develop very slowly. Mature plants white petals, 0.1 to 0.2 inches (2 to 4 mm) in length, will, however, develop in 3 to 4 years. Mountain four stamens, and a two-carpellate pistil. The white to snowberry is easily established using nursery-grown bluish drupes are berrylike, glabrous to pubescent, bareroot or container stock. Seedlings propagated in and 0.28 to 0.35 inches (7 to 9 mm) in diameter. Flesh the nursery or greenhouse grow relatively rapidly, is thin and succulent or mealy. There are usually one producing extensively branched root systems. Soft- or sometimes two bony stones. A smooth rather than wood cuttings taken during the period of flowering or grooved stone is a major feature separating C. s. var. vegetative growth are easily rooted and transplanted stolonifera from C. s. var. occidentalis, although they (Everett and others 1978a; Mirov and Kraebel 1939). are widely hybridized (Hitchcock and others 1961). Redosier dogwood flowers from May to July, depending Uses and Management—This species does not on location. There may be a second period of flowering grow well on disturbed roadways, mine sites, or simi- later in the summer. Fruits ripen from July to Septem- lar disturbances unless areas are top soiled. It has ber and remain on the plant for 8 or 10 weeks (Olsen been seeded and transplanted fairly extensively on and Nagle 1965; Orme and Leege 1980; Stark 1966). such sites with good success. Mountain snowberry is low growing, and is acces- Ecological Relationships and Distribution— sible to all classes of livestock and wildlife. This shrub Redosier dogwood is widely distributed across North is an important source of forage in the early spring, as America, occurring from Alaska to Newfoundland it is one of the first shrub species to leaf out. In areas and south from California to the Northeastern States. where snowberry is a dominant species, it often pro- The species also occurs in central Mexico. It grows at vides important forage throughout the spring and fall. elevations from 1,500 to 10,000 ft (460 to 3,030 m) The persistent, fleshy fruits of all snowberry species (Harrington 1964; Thornburg 1982) on moist sites, are taken by a variety of birds. Field mice scatter the usually along streams, swamps, low meadows, river seeds, and plants are widely established from mice and creek banks, fields, and woods in areas receiving caches. more than 19 inches (48 cm) of annual precipitation. Varieties and Ecotypes—None.

Family Cornaceae ______Cornus stolonifera Redosier dogwood Description—There are about 30 species of dogwood; most occur in the Temperate Zone of the North- ern Hemisphere. Three species occur in the Rocky Mountain region. Redosier dogwood, named for its red to purplish bark is also known as creek dogwood. It is a deciduous, short-lived, rapidly growing, thicket-forming shrub common to moist places (fig. 19). Its spreading clumps of slender stems rise to 15 ft (4.6 m) in height and 20 ft (6 m) in width from a central crown and spreading root system. Stem layers form on decumbent to prostrate stems that simulate stolons. The degree of vegetative spread varies widely. Bark of young branches is bright red to purplish, later turning Figure 19—Redosier dogwood with its distinctive gray green. Lenticels are diamond shaped. Leaves are red bark, opposite leaves, and stoloniferous habit petiolate, opposite, entire, elliptic-ovate to obovate grows in riparian areas and on other moist sites.

630 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

It is one of the earliest shrubby plants to invade bogs 1939). Beaver, bears, and mountain goats use the fruit, and swamps due to the ability of its roots to live wood, and foliage (Murie 1951; Rue 1964). Rabbits, immersed in water (Conway 1949). It is frequently moose, deer, and elk take twigs and exposed winter associated with stands of alder, birch, and willow, and buds. Plants on winter ranges sometimes suffer from in moderately moist areas, with mountain maple, excessive browsing. Kufeld (1973) rated the forage blackberries, or wild roses. Large plants grow singly in value of redosier dogwood as valuable in winter and fall grasslands, incapable of spreading by layering due to and highly valuable in summer. It receives moderate the dense ground cover. Redosier dogwood, highly fall and winter use and heavy summer use from mule adaptable to a range of soil types, is found on sphag- deer (Kufeld and others 1973). The species provides num mats with a pH of 3.2 (Jewel and Brown 1929) and dense summer cover and partial winter cover for birds alkaline soils of pH 8 (Van Dersal 1938). It grows on and other small animals. soils with textures ranging from silty to clayey. In Redosier dogwood is recommended for streambank Wisconsin it is recommended for planting on soils with plantings to stabilize eroding banks and provide shade. a pH of 5.0 to 6.0, base exchange capacity of 6.0 me per It may be used to provide an attractive row in wind- 100 g, and total nitrogen of at least 0.7 percent (Wilde break and conservation plantings in areas with ad- 1946). Roots of redosier dogwood may be associated equate water availability (Cook 1981; Olson and Nagle with vesicular arbuscular mycorrhiza (Barnhill 1981). 1965; Stokes 1977). Varieties selected for ornamental Redosier dogwood will grow in full sun or in shaded use generally have a more compact growth form. In areas. In full sun, the plants are dense and compact landscapes it provides obvious color and a graceful with many lateral branches. In shade, the growth growth form during all seasons (Stokes 1977; Sutton habit is more open and sprawling with few axillary and Johnson 1974). branches, larger leaves, thinner and less curled blades, Plant Culture—Fruits are collected as soon as and lower leaf/total weight ratios. Shrubs growing in they are ripe to reduce losses to birds. Seeds should full sun tend to develop richer purple coloring in the be fully mature when harvested; some planting fall (Sheppard and Pellitt 1976). failures have been attributed to use of immature seeds Use of redosier dogwood in landscaping, along high- (Smithberg 1974). Embryo dormancy is overcome by a ways, and on mine spoils has led to a number of 60- to 90-day wet prechilling at 35 to 41 F (2 to 5 C). physiological studies to determine the effects of short ∞ ∞ A tetrazolium chloride or embryo excision test is nor- days, water stress, defoliation, and low temperature in mally used to indicate seed quality and preclude the initiating and maintaining various degrees of frost lengthy wet prechilling requirement (Heit 1955; hardiness (Chen and Li 1978). Brinkmann 1974d). The double-celled fruits some- Lumis and others (1973) found that of 20 deciduous times contain two viable embryos, which may contrib- shrubs selected from those growing along highways in ute to high germination percentages. Ontario, redosier dogwood exhibited the greatest sen- Field seeding is best accomplished by spot seeding sitivity to aerial drift of deicing salt because its buds untreated seed in fall or wet prechilled seed in spring are exposed in winter. Affected trees exhibited twig on favorable microsites. Seedlings must be protected dieback, a tufted growth pattern, and a greater sus- from competition and browsing. Peterson (1953) found ceptibility of flowering buds than vegetative buds. that laboratory wet prechilling produced better green- Heale and Ormrod (1982) evaluated plants used for house germination than wet prechilling through expo- revegetation of toxic wastes high in nickel and copper sure to winter conditions, or combinations of each type at primary base metal smelters and found redosier of wet prechilling with scarification. Laboratory wet dogwood exhibited typical symptoms. It was sensitive prechilling also provided the greatest resistance to to both levels of nickel tested (2 and 10 mg per 1), but damping off. was sensitive only to the highest copper level tested Nursery seeding is accomplished by seeding freshly (20 mg per 1, but not 4 mg per 1). collected fruits. Dry stored fruits should be soaked in Uses and Management—Redosier dogwood seed- water prior to planting (Heit 1968a). Seeds may also be lings establish and grow rapidly on wet sites, making wet prechilled in the laboratory for spring planting. it a useful species for streambank, erosion control, and Seed should be drilled at a density of 40 viable seeds windbreak plantings. It can withstand moderate brows- per ft2 (430 per m2). Seedlings grow rapidly under ing, but repeated heavy browsing leads to plant loss favorable conditions. One- or two-year-old bareroot (Swihart and Yahner 1983). New plantings begin stock may be used for outplantings (Shaw 1984). producing fruit in 3 to 5 years. Plants may also be grown as containerized seedlings Berries of redosier dogwood provide nutrients for (Landis and Simonich 1984). pheasants, grouse, and many other birds in early win- Redosier dogwood reproduces vegetatively from sto- ter (Stokes 1977). In New England redosier dogwood lons or layering (Peterson 1953; Smithberg 1974). seeds were found in the diets of 95 bird species (McKenny Layering may be induced by pressing branches to the

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 631 Chapter 23 Shrubs of Other Families ground with hooks and covering them with loose soil. Plant Culture—Throughout its natural range, Rooted shoots are later severed, dug, and transplanted. cones develop in late fall and pollen is shed in October Layers, as well as hardwood or softwood cuttings, may and November (Johnson 1974c); seeds mature 15 to 18 be used for vegetative propagation. Dormant cuttings months later. At the time of maturity cones are about may be rooted immediately in the field in moist areas, 1.2 inches (3 cm) diameter. Because cones require or rooted in a nursery or greenhouse prior to field approximately 2 years to mature, numerous cones of planting (Doran 1957). Planting stock is generally different ages are borne on a single tree. Some trees available commercially, but origin should be matched produce abundant female cones; other trees are much as closely as possible to the planting site. less productive. Trees less than 10 years of age normally do not produce many cones. Insects often dam- Varieties and Ecotypes—None. age the cones; consequently, seeds should be carefully inspected to assure healthy cones and viable seeds are Family Cupressaceae ______collected. Ripe cones are dark brown or deep purplish brown (Sudworth 1915; Wolf and Wagener 1948). Cupressus arizonica Seeds are harvested by handpicking or beating the Arizona cypress tree to dislodge the hard woody cones onto canvas tarps. Trees can be damaged from excessive flailing. Description—Arizona cypress is an erect, some- Cones are normally harvested in early to late fall. The what pyramidal, evergreen tree with fragrant, resin- cones are serotinous; they open when mature to shed ous foliage (Johnson 1974c; Welsh and others 1987). the seeds. The scales surrounding the cones form a Five different varieties were described by Johnson tight cluster that can be difficult to open unless quite (1974c); these differ somewhat in erectness and dry. Drying the seeds normally causes natural shat- crown spread (Sudworth 1915; Wolf and Wagener tering in 1 to 2 months. Johnson (1974c) suggested 1948). Trees are mainly 15 to 70 ft (5 to 21 m) in storing the cones for several days in a refrigerator to height (Johnson 1974c; Welsh and others 1987). aid in drying and to prevent cone hardening. Seeds Trunks are 5 to 20 inches (13 to 51 cm) thick; twigs normally separate freely from the cones. Approxi- are stout, branching at nearly right angles; the bark mately 55,050 cleaned seeds per lb (121,400 per kg) is scaly or furrowed and usually grayish. Leaves are were reported by Johnson (1974c) for collections of scalelike, sharply pointed, blue green, and 0.1 inch C. a. variety glabra. Between 90 and 100 seeds are (2 mm) long. Cones are globose, 0.6 to 1 inch (1.5 to produced per cone. Considerably larger numbers are 2.5 cm) thick, short-stalked, hard and woody, with six reported for other varieties (Goggans and Posey 1968; to eight flattened scales bearing a hard point in the Johnson 1974c; Wolf and Wagener 1948). Seeds are center. Seeds are 0.1 inch (2 mm) long and purplish flattened or lens shaped with short wings (Johnson brown (Welsh and others 1987). Plants are monecious. 1974c). Staminate and ovulate strobili are produced on the Seeds wet prechilled for 30 days at 34 ∞F (1.1 ∞C) will ends of the branches. Staminate strobili are long and germinate readily at incubation temperatures near cylindrical, turning yellow as pollen ripens (Johnson 1974c). Ovulate strobili are small, usually less than 0.3 inch (6 mm) long, and green, with six to 12 distinctly arranged scales (Johnson 1974c). Ecological Relationships and Distribution— Arizona cypress is distributed from Texas to Arizona and Mexico (Welsh and others 1987). The varieties described by Johnson (1974c) occur in somewhat different locations, mostly in north-central Arizona, southern California, and Mexico. This plant has commonly been grown as an ornamental and Christmas tree throughout much of the West. It has proven adaptable to areas outside its native range, and has been planted on numerous sites including many range and wildland communities (fig. 20). It is best adapted to sites receiving at least 14 to 16 inches (36 to 41 cm) of annual precipitation. Normal occurrence is on north slopes, coves, benches, and canyon bottoms (Sudworth 1915), Figure 20—A 10-year-old planting of Arizona cy- particularly moist, sheltered canyon bottoms. press provides wildlife cover in south-central Idaho.

632 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

70 ∞F (21.1 ∞C) (Johnson 1974c), but most authors This species has received considerable browsing by recommend incubation at alternating temperatures of big game at all planting sites in the Intermountain 86 ∞F (30.0 ∞C) day, and 68 ∞F (20.0 ∞C) night (Interna- area. Game animals use the plant heavily in summer, tional Seed Testing Association 1966; Stein 1966; and in lesser amounts in the spring, fall, and winter. USDA Forest Service 1948). Seeds may be infested The foliage is quite palatable, and heavy browsing and with mold or bacteria that reduce germination. Treat- highlining of the trees normally occurs. Heavy brows- ment with Captan (N-[(trichloromethyl) thio]-4- ing can prune or eliminate leaf and small stem growth cyclohexene-1,2-dicarborimide) or other fungicides will from the lower portion of the plant. retard fungal development (Johnson 1974c). Because Arizona cypress has demonstrated mixed success on young seedlings are also susceptible to damping off mine sites and other disturbances. It is not adapted to fungi, seedbeds or nursery-grown containers can also infertile mine wastes unless top soiled. It does not be pretreated with a fungicide. serve well as a pioneer species on barren disturbances. Arizona cypress has been established by fall seeding Young plants succumb quickly if planted on margin- in a well-prepared seedbed. Seedlings are sensitive to ally adapted sites. herbaceous competition, and require a weed-free seed- Varieties and Ecotypes—None. bed for 1 to 2 years. Established seedlings are vigorous and grow rapidly when well protected. Plants may attain heights of 2.5 to 4 ft (0.8 to 1.2 m) in 2 to 3 years. Family Cupressaceae ______They continue to grow rapidly, but may not reach mature stature for 40 years. Juniperus horizontalis Seedlings and transplant stock are easily grown, Creeping juniper and the plant is sold by many commercial nurseries. Small stock 6 to 10 inches (15 to 25 cm) high and large Description—Creeping juniper is a native, decum- ball and burlap trees are usually available. bent to procumbent or prostrate shrub that normally attains heights of 8 to 12 inches (20 to 31 cm) (Bifoss Uses and Management—Arizona cypress is an 1947; Great Plains Flora Association 1986; Welsh and important resource in its native range. It provides others 1987) (fig. 21). Plants develop in clumps or mats lumber, wildlife habitat, and watershed protection that may exceed 23 ft (7.0 m) in diameter (Hitchcock (Johnson 1974c). This tree has been particularly use- and others 1969; Stephens 1973). The decumbent or ful for landscape and erosion control plantings in prostrate stems root at the nodes forming well-rooted southern California on a range of soil types at eleva- clumps. Leaves are mostly 0.04 to 0.16 inch (1 to 4 mm) tions below 6,000 ft (1,800 m) (Horton 1949). It is well long, decurrent, opposite, scalelike or awl shaped, and suited as a ground cover or landscape plant because it acute to spinulose tipped. Cones mature the first grows rapidly, is long-lived, and has an attractive season; they are green when immature and ripen to pyramidal growth form (Horton 1949). blue purple or blue black. They are glaucous, 0.2 to Within the Intermountain States, this tree has been used for conservation and wildlife habitat improvement projects. It has been successfully used as a windbreak or cover plant to protect structures and shelter range and farmlands. It has been used to provide cover in mixed plantings with other trees and shrubs. Being evergreen, it is particularly important as winter cover. It provides a dense amount of foliage from the base to the top of the plant. It has survived wind and heavy snowstorms, but some individuals and populations lack cold tolerance to harsh conditions in mountainous regions in the West. This tree species has been useful for wildlife habitat improvement in Utah (Plummer and others 1968). It establishes and grows well, particularly from transplanting in mountain brush, pinyon-juniper, and big sagebrush communities. Seed germination is somewhat erratic, and irregular stands have established from direct seedings. Natural spread by seedling recruitment is poor from all plantings within the Figure 21—Creeping juniper growing as an under- Intermountain area. story with aspen.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 633 Chapter 23 Shrubs of Other Families

0.4 inch (5 to 10 mm) thick, and mostly three- to five- Seeds also require a period of dry afterripening (Pack seeded (Welsh and others 1987). Seeds are reddish 1921). To enhance germination, seeds of most juni- brown and ovoid to spherical (Hitchcock and others pers are subjected to various pretreatments including 1969). Plants are dioecious, and although mature wet prechilling at 40 ∞F (4.4 ∞C) for 30 to 120 days cones may be produced in one season (Welsh and (Barton 1951; Johnsen and Alexander 1974). others 1987), 2 years are normally required (Great When planted in the fall, seed dormancy can usually Plains Flora Association 1986; Miller 1978). be broken by overwinter wet prechilling. Seedbeds should be covered with mulch to ensure the soil re- Ecological Relationships and Distribution— mains moist during the time of germination (Johnsen Creeping juniper is widely distributed in North America and Alexander 1974). Seeds are normally covered to a (Little 1979). It occurs from Montana to northern depth of at least 0.25 inch (6.4 mm) in a firm seedbed Colorado and Utah, and east to northeastern Iowa, (Stoeckeler and Slabaugh 1965). Seedbeds are fre- Nebraska, Illinois, and New York (Little 1979). It is quently shaded during the first growing season to abundant throughout Canada (Couchman and Von protect the soil surface from rapid drying and reduce Rudloff 1965), and occurs from Alaska and the Yukon, the chance of damage from early spring frosts (USDA east to the Atlantic, and south to British Columbia Forest Service 1948). (Welsh and others 1987). The shrub is widely distributed Creeping juniper can be propagated from stem cutover a number of different plant communities. It tings. Portions of the stem that contact the soil and grows in association with grass and broadleaf forbs form roots can be cut and separated into individual at low elevations in the northern Great Plains (Girard plants. The stem cutting and excised root system can 1985; Hansen and others 1984; Johnston 1987). It is be planted in containers, grown to a suitable size, and also associated with conifer forests, aspen, and limber field planted. Stem cuttings can also be rooted easily pine (Girard 1985) at high elevations and on more under greenhouse conditions (Zorg 1954). Stem cut- moist sites (fig. 21). tings taken in early winter (November and December) Plant Culture—Staminate cones are produced at root readily without the use of rooting compounds the tips of the branches in early spring. Pollination of (Doran 1957; Kiplinger 1938). pistillate cones normally occurs in April and May Plantings of 2-year-old bareroot stock have estab- (Miller 1978). Pistillate cones form globose, berrylike lished and survived better than younger, smaller stock. fruits (Great Plains Flora Association 1986; Hitchcock Field lifting and transplanting must be completed in and others 1969; Miller 1978). As they ripen they early spring when plants are dormant. Although trans- change in color from green to blue black or bluish planting may be done later in the season, supplemen- purple (Stephens 1973). Cones normally remain on tal irrigation is usually required to assure survival. the plant until the late fall or early winter of the second year. When fully mature they drop from the plant. Uses and Management—Creeping juniper has Seeds are harvested by stripping or picking the not been used extensively in large revegetation projects. ripened cones. Cones can also be dislodged by flailing Although recognized as an important ground cover the branches and dislodging them onto canvases spread and a locally useful forage plant for wildlife, it has on the ground. It is advisable to collect the cones as been difficult to establish on wildland sites. This soon as they ripen. Collecting green cones or immature species has been able to persist following heavy graz- seeds is not advised, as they are difficult to separate ing when more palatable shrubs and herbs have been from fully ripened seeds during cleaning. Freshly eliminated. Consequently, rehabilitation programs collected cones should be spread to dry, but can be often do not include this shrub. Creeping juniper stored and seeded as dry cones or as cleaned seeds normally occurs as scattered plants, occupying ridge (Johnsen and Alexander 1974). Seeds are normally crests, rocky slopes, ledges, washes, or areas that are extracted from the cones by maceration using a Dybvig difficult to plant with conventional equipment. Al- seed cleaner. The debris is floated away in water though site accessibility limits planting in some areas, (Johnsen 1959; Johnsen and Alexander 1974). If cones this plant is important for ground cover and wildlife are completely dry before cleaning, they may require habitat and should not be overlooked. soaking for a few days to aid in separation. The Creeping juniper provides dense cover on harsh macerated material is dried and then fanned to sepa- sites. Its low evergreen growth habit provides diver- rate seeds from the trash. Seeds should be stored in sity and cover throughout the entire year. Ecotypes that are semierect or upright furnish better habitat for sealed containers at 20 to 40 ∞F (–6.7 to 4.4 ∞C) with relative humidity at about 10 percent (Johnsen and wildlife. This shrub grows on sites where few other Alexander 1974; Jones 1962). plants are adapted. Because creeping juniper grows on Germination of creeping juniper seed is delayed by isolated wildlife ranges, it is important to maintain its embryo dormancy and an impermeable seedcoat. presence. It occurs in most plant communities and big game winter ranges on sites where soil protection and

634 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families forage diversity are critical for protecting watershed occurs. Use of this shrub has been encouraged where and wildlife values. Consequently, it is important that efforts are made to restore native communities. Creep- techniques are developed to better accommodate its ing juniper has established slowly on mine spoils, and propagation and field planting. appears sensitive to soil conditions. Although it natu- Creeping juniper normally receives only limited use rally occurs on infertile sites, it is not well suited to all by big game, but locally heavy winter browsing has mine wastes and spoil materials. It does best on sites been reported (Frischknecht 1975; Miller 1978). Dusek that have been top soiled or contain soil fines capable (1975) found that creeping juniper provided a major of holding soil moisture. It has been less successful on portion of mule deer diets during March and April in rocky, coarse mine wastes and exposed substrata. To north-central Montana. This plant provides impor- be successful on mine disturbances, a balanced amount tant fall, winter, and spring forage for big game (Dusek of soil nutrients must be provided. 1971; Kufeld and others 1973; Lovaas 1958; Martinka This shrub can be used to restore mine spoils where 1968; McKean 1954; Schallenberger 1966). It is more irregular topography is created. If topsoil or surface commonly used by deer, but Dirschl (1963) found that soil material is reapplied, the plant can be successfully creeping juniper provides about 10 percent of the planted to re-create natural communities and land- winter diet of pronghorns, and was the dominant scapes-disturbed areas to match adjacent native forage from December to March in some parts of plant associations. Saskatchewan. Some ecotypes in the Intermountain Creeping juniper can also be used to stabilize water- States exhibit considerable differences in use by game sheds and erodible sites. It establishes and grows animals. Selections that receive moderate or heavy slowly, a major disadvantage in stabilizing erosive use are consistently browsed at these levels, although soils and steep slopes. However, once established, it is climatic, edaphic, or other factors may contribute to able to withstand windy conditions, and provides selective use by game. excellent ground cover under very exposed and harsh Birds and other wildlife seek sites occupied by situations. This shrub is also very useful as a land- this shrub. The cones provide food for many game scape species. It is widely used in horticultural birds and mammals. They remain on the shrub for 1 or plantings in areas including recreational sites, camp- 2 years and provide 20 to 40 percent of the total food grounds, parks, and associated sites where little main- consumed by sharptail grouse (Miller 1978). Not all tenance is provided. It is particularly useful in plant- plants produce heavy seed crops. Annual production is ing parks and recreational sites where use of native often quite low, but fruits that do develop are used species is emphasized. It can withstand heavy traffic, during winter and early spring periods when other and is used to align walkways, roadways, and trails. foods are less available. Varieties and Ecotypes—Various selections have Creeping juniper begins growth very early in spring. been developed for horticultural uses that exhibit Game animals are attracted to the shrub in late different colors, or vary in growth habit, leaf texture, winter and early spring when other forages are less rooting habits, and growth rates. However, none have palatable. The plant grows in areas where game may been released for wildland situations. be forced to concentrate during periods of deep snow and adverse weather conditions. At these critical periods, it provides protection, cover, and forage that Family Cupressaceae ______are crucial for the survival of game animals. Creeping juniper has been used to restore disturbed Juniperus osteosperma watersheds and mine sites. It is extremely difficult to Utah juniper establish by direct seeding because of germination problems. Transplanting has been the most successful Description—Utah juniper is a small tree nor- means of establishment; however, costs associated mally with a single stem and several short branches with transplanting limit its usefulness. Transplants that originate near the ground (fig. 22). Individual are difficult to establish on semiarid sites. Small trees may exceed 21 ft (6.4 m) in height (Goodrich and transplants root slowly and require frequent irriga- Neese 1986). They have thin, fibrous bark that shreds tion to fully establish. Thus, it has been difficult to from the trunk and branches (Welsh and others 1987). plant sites that do not remain moist through the year Welsh and others (1987) described the mature leaves of planting. Once established and well rooted, this as typically opposite, scalelike, 0.04 to 0.12 inch (1 to species survives very well. New plantings are not able 3 mm) long, and yellowish green. Juvenile leaves are to compete well with broadleaf herbs and grasses, and decurrent, awl-shaped, sharp, and 0.1 to 0.3 inch (2 to the shrub should be planted alone. 8 mm) long. Plants are monoecious; staminate cones This shrub occupies areas in Wyoming, Montana, are yellowish brown and 0.1 to 0.2 inch (3 to 4 mm) and Colorado where considerable surface mining long; ovulate cones are subglobose, 0.2 to 0.5 inch (6 to

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 635 Chapter 23 Shrubs of Other Families

concluded that the distribution of Utah juniper is primarily regulated by precipitation. Tree stands normally occupy sites that receive from 10 to 25 inches (254 to 635 cm) of annual rainfall (Blackburn and Tueller 1970). Utah juniper is more tolerant of fire and spring frost, and performs better with low summer precipitation than singleleaf pinyon pine (Tueller and others 1979; West and others 1979), yet these factors do not fully explain the distribution and composition of the tree species and communities. At higher elevations, singleleaf pinyon may dominate and is generally more abundant. Utah juniper is widely adapted to a variety of soil types and parent materials. It grows on limestone, sandstone, granite, and mixed materials (Clary and Figure 22—Stands of Utah juniper furnish consid- Jameson 1981). Utah juniper is particularly well erable cover for a variety of wildlife following adapted to rocky, well-drained, coarse-textured soils chaining and seeding with grasses and forbs. (Eckert 1957). Plant Culture—Good seed crops are normally produced annually by most wildland stands, although abundant seed crops may only occur every 2 to 6 years. Seeds require about 2 years to mature. Cones appear 12 mm) thick and brownish or blue to blue purple at from mid-April to mid-May, and normally mature in maturity. There are one to two seeds per cone (Arnold late fall or early winter the second year after flowering and others 1964; Johnsen and Alexander 1974; Tueller (Johnsen and Alexander 1974). Cones of different ages and Clark 1975; Welsh and others 1987). are normally found on the same plant. Mature cones Ecological Relationships and Distribution— are darker in color and are obviously drier than imma- Utah juniper occurs from Montana and Wyoming, ture cones (Zarn 1977). south to California, Arizona, and New Mexico (Griffin Most wildland stands produce commercially and Critchfield 1972; Hitchcock and Cronquist 1973; harvestable crops, although selected individuals or Welsh and others 1987; Zarn 1977). It grows with small groups of trees may be the primary producers. singleleaf pinyon to form vast woodlands, typically Seed collection, cleaning, and storage practices are existing between more xeric cold desert shrub com- similar to those described for creeping juniper and munities at lower elevations and mountain brush or discussed by Johnsen and Alexander (1974). ponderosa pine at higher elevations. It normally occurs As with other junipers, the seed coats are imperme- between elevations of 6,000 to 7,500 ft (1,800 to 2,300 m) able and embryos are dormant (Johnsen 1959; Johnsen (Goodrich and Neese 1986), although it is not uncom- and Alexander 1974). Seeds are difficult to germinate mon between elevations of 2,790 to 8,000 ft (850 to without pretreatment. Johnsen and Alexander (1974) 2,400 m) (Welsh and others 1987). recommended a 120-day warm pretreatment at 68 to Utah juniper is the dominant species, along with a 86 ∞F (20 to 30 ∞C) alternating diurnal temperatures number of shrubs and grasses, on many sites that have followed by wet prechilling for 120 days at 41 ∞F (5.0 ∞C). been subjected to heavy domestic grazing (Clary 1975a) Incubation at alternating day and night temperatures and reduction of wildfires. These management prac- of 68 to 86 ∞F (20.0 to 30.0 ∞C) for 70 days resulted in tices have resulted in an increase of trees, coupled less than 50 percent germination. with a decrease of understory plants. Pinyon-juniper Although Utah juniper spreads well by natural stands have not only increased in density, but have seeding, it is a difficult plant to propagate and it spread to occupy adjacent shrub and grass communi- establishes poorly by artificial seeding. Field plantings ties (West and others 1979). The spread of Utah establish erratically and unpredictably. Erratic seed juniper may be regulated by periods of drought and the germination is the primary factor affecting poor seed- occurrence of years of favorable precipitation ing establishment. Pretreatments to stimulate and (Blackburn and Tueller 1970). regulate seed germination have been somewhat help- Utah juniper is more tolerant of arid regions in the ful when seeds are germinated under controlled labo- Great Basin (Tausch and others 1981) than singleleaf ratory conditions. However, pretreatment practices pinyon; it is the only tree species present in many have not significantly improved seedling emergence areas. It forms dominant stands throughout much of from wildland plantings. Seedling establishment in southern Utah and Nevada. Treshow and Allan (1979) nurseries has been enhanced by prechilling seeds or by

636 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families mulching and irrigating the seedbeds, but erratic 2- to 3-year-old stock has exceeded that of smaller stands still result (Johnsen and Alexander 1974). transplants. Small transplants grow slowly and do not Seedlings of Utah juniper that emerge in early compete as well as larger stock. Utah juniper trans- spring and survive spring frosts are most likely to plants do not survive well if the primary root of young survive their first season. Seedlings that germinate plants is damaged. This impact is less evident for late in spring, as soil moisture is depleted, are less larger stock. likely to survive. New seedlings grow slowly and Utah juniper can be propagated from stem cuttings normally require 2 to 4 years to attain large stature. taken from November through February, but it is During this period of establishment, mortality is very difficult to root (Doran 1957; Snyder 1954). Treatment high. Young plants that do survive for 1 to 4 years with root-inducing substances may or may not be become hardy, and can persist with considerable her- beneficial (Doran 1957; Zorg 1954). baceous competition and adverse climatic conditions. Uses and Management—Utah juniper is widely Blackburn and Tueller (1970) concluded that spread of regarded as an important plant for big game winter juniper is favored by years of good seed production ranges. The plant provides important cover and for- followed by 6 years of average or above-average pre- age for wildlife (Frischknecht 1975). Nearly all big cipitation. Seedling establishment is difficult to pre- game animals utilize pinyon-juniper woodlands. dict. Numerous chaining and burning projects have This species is not regarded as highly palatable to big cleared areas and created openings where new seed- game by most authors (Brotherson and Osayande lings may establish. Chaining and other activities 1980). Rosenstock and others (1989) found that mule bury surface-deposited seeds, but new tree seedlings deer depended heavily on juniper during winter do not always invade the treated sites. New tree months, and it is a principal part of their diet during seedlings may not appear for many years after treat- the midwinter period. The species may be regarded as ment, and increases may occur after understory spe- an emergency winter forage plant (Gullion 1964), cies have established a dense cover. with only fair energy and protein values (Dittberner Tree seedling survival is affected by the presence and Olsen 1983), but Kufeld and others (1973) con- and density of understory herbs. Monsen and others cluded that juniper is moderately to heavily grazed by (1987) found that native understory species, including big game during most seasons. The fruits are widely shrubs, have prevented tree seedling encroachment in used by numerous birds (Turkowski and Watkins central Utah. Seeding introduced grasses and broad- 1976). Big game and livestock graze the understory of leaf herbs on pinyon-juniper sites reduces the estab- pinyon-juniper sites heavily in spring, fall, and winter. lishment of tree seedlings, but the influence of indi- Utah juniper provides essential cover for big game, vidual herbaceous species is not fully understood. The particularly during the winter months. Animals seek presence of native herbs also limits tree seedling cover and protection, and current chaining practices survival. are now being carefully evaluated to assure that ad- Tree recruitment is undoubtedly site influenced. equate sized openings, escapeways, and concealment Natural recruitment frequently occurs very rapidly on cover are retained. both treated and untreated sites where mature trees Game animals, livestock grazing, and recreational have been removed by burning or chaining. On adja- uses of pinyon-juniper sites have increased dramati- cent treated areas, tree recovery has been delayed for cally in many locations. Game animals have been extended periods even though all sites receive similar forced to use pinyon-juniper areas in the spring, fall, amounts of precipitation. In addition, at some Utah and winter, as access to and use of natural ranges have locations, Utah juniper stands cleared of mature trees been diminished by the construction of homes, roads, at different times, have little tree recruitment, but farms, ranches, and commercial developments. Many other sites have experienced considerable tree recov- pinyon-juniper sites have been converted to shrub/ ery each year after treatment. grass communities and managed to prevent tree re- When direct seeded, Utah juniper should be planted cruitment. Pinyon and juniper have also spread to at a depth of 0.5 to 1.0 inch (1.3 to 2.5 cm) and sown occupy sites that are void of competitive understory in late summer or fall (Heit 1967b; Johnsen and species. The encroachment has been facilitated by Alexander 1974; Stoecker and Slabaugh 1965). Hand the reduction or control of wildfires. Consequently, seeding in small openings where the soil surface is pinyon-juniper sites have become increasingly im- protected by litter, rock, or other debris tends to portant to the management of wildlife populations. improve survival. Chaining, burning, and seeding of pinyon-juniper Utah juniper is widely used in landscape plantings areas for livestock grazing may not be economically as 2- to 5-year-old container transplants (Johnsen practical in all situations, but restoration and improve- and Alexander 1974). Bareroot transplants have also ment for wildlife habitat improvement is beneficial. been cultivated for field plantings. Survival of larger

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 637 Chapter 23 Shrubs of Other Families

Pinyon and juniper communities are an important Family Cupressaceae ______component of many watersheds. These sites may not receive great amounts of precipitation each year, but Juniperus scopulorum plants grow on steep slopes that are subjected to runoff Rocky Mountain juniper and erosion from high-intensity storms. Closed stands of pinyon-juniper and communities that lack suitable Description—Rocky Mountain juniper is typically understory provide poor ground cover and are high taller than Utah juniper, normally reaching 10 to sediment producers. These sites should be managed to 20 ft (3.0 to 6.1 m) in height (Welsh and others 1987), prevent infrequent, but damaging erosion and runoff but heights of 20 to 50 ft (6.1 to 15.2 m) are not from unpredictable storm events. Pinyon-juniper wood- uncommon (Johnsen and Alexander 1974). Crowns lands are also subjected to wildfires and other impacts are conical to pyramidal or, less commonly, rounded. that can diminish tree cover. If the tree communities The bark is thin, fibrous, and shredded. Leaves are lack a suitable understory, the burned or disturbed opposite, but sometimes in threes, scalelike, 0.02 to sites become vulnerable to erosion; this significantly 1.2 inches (0.5 to 30 mm) long, green or blue green, impacts downstream resources. Consequently, it is sometimes with dorsal resin glands. Juvenile leaves important to maintain a diverse understory of shrubs are needlelike and 0.1 to 0.3 inch (3 to 8 mm) long and herbs that are capable of recovering following (Welsh and others 1987). Plants are generally monoe- these natural disturbances. cious with staminate cones 0.08 to 0.12 inch (2 to 3 mm) Pinyon and juniper communities have been invaded long; female cones develop strobili or berrylike struc- by cheatgrass, even where closed tree stands exist. As tures that are 0.2 to 0.3 inch (4 to 8 mm) thick at mature trees die from fires, insects, or disease, the maturity (Johnsen and Alexander 1974; Zarn 1977). annual grass spreads quickly, and once in place pre- Arnow and others (1980) distinguished Rocky Moun- vents the recovery of nearly all native species. Exten- tain juniper from Utah juniper by leaf shape; Rocky sive areas of pinyon-juniper woodlands throughout Mountain juniper has entire and opposite leaves rather the Intermountain region occur with only remnant than minutely toothed and somewhat whorled leaves. amounts of native shrubs and herbs, but with scat- Ecological Relationships and Distribution— tered and suppressed amount of cheatgrass. As these Rocky Mountain juniper occurs throughout central sites eventually burn, the sites are converted to an- British Columbia and Alberta, and much of the Pacific nual grass communities. The annual grass ranges Northwest, Rocky Mountains, and Great Basin. It is burn frequently, resulting in a loss of wildlife habitat. found from the northern Great Plains, south to Ne- To prevent the conversion of pinyon-juniper sites to vada, Arizona, New Mexico, and Texas (Fowells 1965; annual grass, sites can be artificially seeded with Hitchcock and Cronquist 1973; Pieper 1983; Welsh desirable understory natives following burning or and others 1987; Zarn 1977). It is commonly encoun- other disturbances. In addition, potential problem tered in canyons and cool exposures at higher eleva- areas can be treated by controlled burns or chaining tions than Utah juniper, normally occurring as scat- followed by seeding. This practice has been exten- tered plants rather than extensive stands (Goodrich sively employed in Utah and other Intermountain and Neese 1986). It usually grows with mountain big States. This problem has not been fully understood sagebrush, mountain brush, ponderosa pine, aspen, and and addressed in land management programs. Douglas-fir at elevations between 7,000 and 9,000 ft Deterioration of pinyon-juniper communities within (2,100 and 2,700 m) (Fowells 1965; Goodrich and the Intermountain area has had a significant impact Neese 1986). At higher elevations it grows with on big game populations. Most pinyon-juniper sites Engelmann spruce and subalpine fir. Although a have been heavily impacted by previous grazing by typical mountain species, it often occurs at low eleva- livestock and wildlife. This has resulted in a loss of tions in valley bottoms with cold air drainage (Welsh shrub and herb understory. Trees have increased and and others 1987). It is most common in open wood- once in place prevent the return of the desired under- lands or with big sagebrush-grass communities. story. Consequently, extensive chaining, clearing, and Throughout central Utah it is particularly abundant seeding projects have been employed to reduce tree with Gambel oak. The erect columnar variety is more density and allow seeded species to establish. Numer- common in Utah on open aspects with mountain big ous native shrubs, broadleaf herbs, and grasses have sagebrush. It is also common in northern Wyoming been developed to convert tree-dominated sites into and southern Montana. sites dominated by a more diverse array of species. Plant Culture—Cones appear from mid-April to mid- Varieties and Ecotypes—None. June, mature from mid-September to mid-December

638 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families the second year after pollination, and persist on the evergreen growth habit provides attractive yearlong plant for 2 to 3 years (Fowells 1965; Johnsen and protection. The plant is sufficiently drought tolerant Alexander 1974). Seeds are harvested, cleaned, and to persist on sites receiving 12 to 14 inches (30.5 to 35.6 stored as described for other junipers (Johnsen and cm) of precipitation. It is better adapted to alkaline soil Alexander 1974). Seed production is erratic, but some conditions than most other junipers (Miller and others seeds are produced every year; abundant crops are 1948). Rocky Mountain juniper is particularly useful produced about every 2 to 5 years. Seeds are collected when planted in combination with other trees and by hand stripping or beating the limbs to dislodge the shrubs as a windbreak or screen. It withstands harsh, cones onto canvas spread on the ground. Seeds are cold temperatures, and can be used to protect road- relatively small compared with those of other juni- ways, buildings, and other facilities. pers; the count ranges between 17,850 and 42,100 This species is also of importance as a cover and seeds per lb (39,350 and 92,800/kg) (Johnsen and forage plant for big game. It receives moderate use in Alexander 1974). winter, spring, and fall (Kufeld and others 1973). Germination is delayed because of embryo dormancy However, considerable differences are noted among and hard impermeable seedcoats; seed pretreatments sites and game animals. Some plants are grazed are recommended to hasten germination. Untreated heavily, but most trees are not grazed extensively. seeds may require 14 to 16 months to germinate Livestock generally make little use of this plant. (Afanasiev and Cress 1942). Johnsen and Alexander Rocky Mountain juniper furnishes excellent cover (1974) reported that Rocky Mountain juniper seeds for a variety of wildlife. It frequently grows in associa- require a warm pretreatment at 68 ∞F (20 ∞C) for 45 to tion with various shrubs that provide winter cover. Big 90 days followed by wet prechilling to initiate game animals often use stands of Rocky Mountain germination. juniper for cover and protection. Sites that are used for Johnsen and Alexander (1987) suggested different this purpose should be managed to maintain appropri- methods of pretreatment and seeding. These include: ate cover and habitat. (1) cleaning the seed followed by warm/cold wet pre- Rocky Mountain juniper frequently grows in canyon treatment and fall seeding; (2) storage of the cones bottoms, streams, seeps, and other riparian areas. It (berries) for 1 year before cleaning, wet prechilling, provides excellent cover and structural diversity. Plants and fall or spring seeding; and (3) a warm pretreament also normally support a herbaceous understory if sites outdoors from spring to the time of fall sowing. are properly managed. Small pockets of Rocky Moun- Seeds should be planted in a firm seedbed at a depth tain juniper, intermixed with mountain big sagebrush, of 0.25 to 0.5 inch (6 to 13 mm). Addition of a surface are important areas for wildlife. The diversity pro- mulch is helpful. When planting wildland sites, spots vided by this plant is important for many upland sites should be selected that are somewhat protected and where lower growing shrubs occur. have variable soil conditions. Seedlings do not emerge Few other tree species are able to grow where this from bare, open, compacted, or crusted surfaces. native normally exists. Because it is relatively slow to Young Rocky Mountain juniper plants normally establish and mature, removal of this species should grow faster than Utah juniper until they reach mature be avoided. Restoration projects should be designed to stature. Fowells (1965) reported fairly uniform growth to an age of 40 years, at which time the growth rate declines. Plants live 250 to 300 years (Tueller and Clark 1975). This species is easier to rear as bareroot or container stock than Utah juniper. Seeds are somewhat less dormant, and plants usually grow faster when irrigated and fertilized. Commercial growers often produce this species and are able to rear the plant with good success. Transplant stock of various ages is produced and sold as potted container stock or ball- and-burlap plants. Plants are usually 3 to 4 years of age before they are of sufficient size for commercial sales. Uses and Management—Rocky Mountain juniper (fig. 23) is commonly used for conservation, windbreak, and horticultural purposes. This species has been useful for improving upland game bird habitat on Figure 23—Young Rocky Mountain juniper trees farmlands (Miller and others 1948). Its pyramidal grow in association with mountain big sagebrush.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 639 Chapter 23 Shrubs of Other Families leave this species in areas of high importance. Rocky Mountain juniper normally grows on sites that are moist enough to support understory herbs. Herb and shrub seedlings are better able to establish with this overstory tree than with Utah juniper. Direct seeding of herbs can be recommended on areas where scattered plants exist. Complete removal of the trees is not required to accomplish seedling establishment of desired native understory species. Rocky Mountain juniper has been planted on mine disturbances with mixed success. Small 1- to 2-year- old container stock has not established or persisted well on mine wastes in southeastern Idaho and southern Montana. Larger stock, 2- to 6-year-old potted plants, survive better than smaller stock. Supplemen- Figure 24—Silverberry grows along stream bottoms tal irrigation for 2 to 3 years significantly improves with willows and other riparian species in eastern Utah. establishment on harsh sites. Once plants are well rooted, they persist well. Varieties and Ecotypes—None. Plant Culture—Silverberry plants flower in early Family Elaeagnaceae ______summer. The large, silver-green drupelike fruits ripen in late fall and early winter. This shrub produces Elaeagnus commutata abundant seed crops at intervals of 1 to 2 years. Ripe Silverberry fruits can be picked by hand, or the bushes can be beaten to dislodge the fruits onto canvas spread on the Description—Silverberry is a native, deciduous ground (Olson 1974). The fruits are easier to clean shrub, mostly 3 to 6.5 ft (0.9 to 2.0 m) tall, with peltate before they dry. Seeds are extracted from the fruit by scales. Plants vary in growth habit, often occurring as maceration in a Dybvig separator, and water is used to thicket-forming shrubs up to 12 ft (3.7 m) tall or grind and float the pulp away (Heit 1968a; Olson frequently as small trees (Johnson and Anderson 1974). Between 2,700 and 4,600 cleaned seeds are 1980). Stems are stoloniferous and unarmed. Leaves reported per lb (5,950 to 10,140 per kg) of pure seed are 0.5 to 2.8 inches (1.3 to 7.1 cm long), 0.2 to 1.2 (Olson 1974). inches (6 to 31 mm) wide, elliptic to oblanceolate, Silverberry has not been widely used in range and acute to obtuse or rounded and silvery on both sides wildlife projects. Seeds should be planted 0.5 to 1 inch with brown scales. Flowers are one to four per axil, (1.3 to 2.5 cm) deep in a firm seedbed. Seedling growth 0.4 to 0.6 inch (1.0 to 1.5 cm) long, and yellowish. The rate is fairly rapid, and young plants persist quite well fruit is drupelike, consisting of a dry indehiscent if planted in adapted areas. Plants also establish well achene enveloped by a fleshy persistent hypanthium. from bareroot or container-grown stock. Silverberry is Fruits are 0.2 to 0.4 inch (6 to 10 mm) long, mealy, and reported to have stoloniferous stems (Johnson and silvery (Welsh and others 1987). Anderson 1980), and can spread from underground Ecological Relationships and Distribution— rootstocks (Viereck and Little 1972). Vegetative spread Silverberry occurs from Quebec to the Yukon, and has not occurred in wildlife plantings in Utah. south to Minnesota, Wyoming, Utah, and northern Uses and Management—Silverberry has been se- Colorado (Harrington 1964). In Utah, it is usually lectively used throughout the Western United States. restricted to riparian zones at elevations between It was propagated as early as 1813 (Olson 1974) for 6,000 and 8,000 ft (1,800 and 2,400 m) (Welsh and windbreaks and shrub row plantings, but has not others 1987). Plants occur in areas receiving 18 to 20 been used extensively in recent programs. The spe- inches (45.7 to 50.8 cm) of annual precipitation, but cies has been recommended for windbreak and wild- usually require supplemental water even on moist life plantings in Wyoming (Johnson and Anderson sites (fig. 24). Silverberry is very winter hardy and 1980). Plants are grown as ornamentals and wind- drought tolerant. In eastern Utah, silverberry grows breaks in the interior of Alaska (Viereck and Little along streams and dry hillsides (Goodrich and Neese 1972). Plants produce leaves having an unusual sil- 1986). In the interior of Alaska it grows on rocky, ver-green cast. Leaves persist on the shrub until south-facing slopes and sandbars (Viereck and Little early winter, thus furnishing an attractive back- 1972), often forming thickets along major streams. ground screen.

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In Utah, big game have made moderate use of the Ecological Relationships and Distribution—Sil- shrub during the spring and fall months, but they ver buffaloberry is distributed from British Columbia seem to prefer the plant much more as summer to Saskatchewan, and south to California, Nevada, browse. Reports by Kufeld and others (1973) and New Mexico, and North Dakota. It occurs along Dusek (1971) described limited fall grazing but in- streams in sagebrush and pinyon-juniper woodlands creased summer browsing by big game. The shrub has (Sampson and Jesperson 1963). It is widely scattered been transplanted with good success on mine distur- throughout Utah at elevations between 4,590 and bances in southeastern Idaho, and has received only 7,510 ft (1,400 and 2,290 m) along streambanks, ter- limited use by mule deer. races, and on moist or wet sites (Welsh and others Silverberry appears to have particular value for 1987). It occupies ditch banks, abandoned fields, dis- riparian habitat improvement within its native range. turbed fence lines and powerlines (Goodrich and Neese It transplants very well and grows rapidly even with 1986). In Wyoming it grows in bottomlands, on moist some understory competition. It is able to establish hillsides, and on streambanks at elevations up to and grow on moist and well-drained soils, typical of 7,000 ft (2,100 m) (Johnson and Anderson 1980). disturbed riparian sites. It provides considerable cover Plant Culture—Plants flower in spring, beginning for animals and soil protection. It is a well-rooted plant in early April and continuing into June. Fruits ripen that can provide excellent soil stability, but it does not from June to August, and they remain on the bush spread as does Russian-olive to become a serious weed. until December (Plummer and others 1968; Thilenius Silverberry has considerable value as an ornamen- and others 1974a; Van Dersal 1938). Ripe fruits are tal plant for residential and recreational sites. It can most easily collected by flailing the plant, causing the be used to enhance parkways, campgrounds, and rec- fruits to drop onto canvas spread on the ground. Fruits reational sites, particularly near streams and semiwet may be handpicked, but the thorny branches hinder areas. It can be used to screen and protect structures, this approach. roadways, trails, and campgrounds. It does not attract Stems, twigs, and leaves must be separated from the heavy use by wildlife and can be used where animal fruits before they are macerated and the pulp floated concentration is to be avoided. away with water. The seeds (cleaned achenes) are Varieties and Ecotypes—None. processed with a fanning mill to separate them from small debris. Seedlots can be cleaned to a high purity. There are between 18,000 and 67,000 seeds per lb Family Elaeagnaceae ______(39,700 and 147,700 per kg); the average is 41,000 Shepherdia argenta seeds per lb (90,390 per kg) (Thilenius and others Silver buffaloberry 1974a). Mirov and Kraebel 1939 and Plummer and others (1968) reported the average number of cleaned Description—Silver buffaloberry is a deciduous, seeds per lb is between 10,800 and 18,000 (23,810 and thorny shrub with a spreading to ascending growth 39,680 per kg). Cleaned seeds can be stored in open habitat that attains heights of 6 to 10 ft (1.8 to 3.0 m) warehouses for at least 5 years without appreciable (Thilenius and others 1974a; Welsh and others 1987). loss of viability (Plummer and others 1968). Goodrich and Neese (1986) reported that plants may Both embryo dormancy and hard seedcoats restrict reach heights exceeding 22 ft (6.7 m). Branchlets are or delay germination. Thilenius and others (1974a) covered with silvery peltate scales and terminate in found that wet prechilling for 60 to 90 days, followed sharp thorns. Leaves are opposite, petiolate, 0.2 to 2.4 by exposure to alternating day and night tempera- inches (0.5 to 6 cm) long, and 0.12 to 0.55 inch (3 to 14 tures from 86 to 69 ∞F (30.0 to 20.0 ∞C) for 30 to 60 days, mm) wide. They are oblong, elliptic or lanceolate, and resulted in 26 to 93 percent germination. silver gray on both sides. Plants are dioecious, with Seeds sown in fall will germinate in spring. Seeds axillary flowers that can be perfect or imperfect. Flow- should be planted at a depth of 0.5 to 1 inch (1.3 to ers are 0.1 to 0.16 inch (2.5 to 4 mm) long with 2.5 cm). Seedlings and young plants have established yellowish perianths. The red fruit is drupelike, 0.16 to moderately well on range or conservation plantings. 0.28 inch (4 to 7 mm) long, and edible, though tart Seedling growth is only fair, but as plants reach 1 or 2 (Goodrich and Neese 1986; Sampson and Jespersen years of age, they become well established and are very 1963; Welsh and others 1987). Staminate flowers have persistent. Seeds are easily handled and planted with eight stamens alternating with glandular thickenings most conventional planters. Bareroot or container- at the base of the perianth lobes (Welsh and others grown transplants establish better than direct 1987). The perianth of pistillate flowers is short and seedings. The shrub is not browsed very heavily, nor tubular, investing the ovary. adversely affected by insects, consequently it is able to establish without serious delays (Plummer and others

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 641 Chapter 23 Shrubs of Other Families

1968). Silver buffaloberry grows moderately well with herbaceous understory species. If 1-year-old transplant stock is planted, very good survival and excellent growth can be attained. The shrub is produced by many commercial nurseries. Root sprouts can be propagated with good success. Uses and Management—Silver buffaloberry has been widely planted for shelterbelts, game habitat, cover, and watershed protection (Thilenius and others 1974a). It has been used in ornamental plantings because of its gray, willowlike foliage (Sampson and Jespersen 1963). The thorny nature of this plant limits its use in heavily populated areas. However, it can be planted in recreation sites to screen areas, direct foot traffic, and control or restrict use of camp- Figure 25—Silver buffaloberry has been successfully grounds and parks. It is used as the outer row in established on mine disturbances in southeastern multirow shelterbelt plantings, but it is often planted Idaho. alone where a low, dense cover is required (Johnson and Anderson 1980). The shrub is highly susceptible to heart rot, and breakage by wind and snow results as the wood becomes brittle (George 1953). In the Mid- overstory cover, thickets, and low ground cover. It is west, high winds have uprooted cultivated trees; con- adapted to streambanks, sandbars, unstable soils, sequently, this species is not recommended for wind- and moist sites in pinyon-juniper and sagebrush break plantings, particularly near buildings and communities where few upright species grow. It trans- walkways. (George 1953). plants very well and can be used to populate open, Silver buffaloberry is recommended for use in pinyon- exposed sites. juniper, big sagebrush, inland saltgrass, and wet Varieties and Ecotypes—None. meadow communities of Utah for wildlife habitat improvement (Plummer and others 1968). It is especially useful on sandy soils and in moist sites Family Elaeagnaceae ______(Thornburg 1982). The thorny thickets created by this shrub provide cover and nesting sites for birds and Shepherdia canadensis other animals; quail (Martin and others 1951) and Russet buffaloberry, soapberry some small mammals take the drupelike berries Description—Russet buffaloberry is a native, thorn- (Hall 1946). Big game animals make considerable less, small to medium, spreading shrub 3 to 9 ft (0.9 to use of the shrub during the summer but only moder- 2.7 m) tall (Thilenius and others 1974a). Branches are ate use in the winter (Compton 1966; Dusek 1971; covered with brown peltate scales. Leaves vary in Kufeld and others 1973; Lamb 1968; McKean 1954; length from slightly less than 0.25 to 3 inches (0.5 to Plummer and others 1968). 8 cm). They are ovate to lanceolate, rounded apically Although silver buffaloberry is normally recom- and basally, green above and slightly pale beneath. mended for moist site plantings, it has performed well Flowers vary from 1 to several per axil, and 0.08 to 0.12 on mine disturbances in southeastern Idaho, Mon- inch (2 to 3 mm) long. Fruits are 0.16 to 0.28 inch (4 tana, and Wyoming (fig. 25). It has established and to 7 mm) long, red, and quite succulent (fig. 26). When grown well on sites receiving 16 to 24 inches (40.6 to mixed with water the fruits produce a soapy solution 61.0 cm) of annual precipitation. The species has (Welsh and others 1987). established from transplant stock on mixed, infertile mine wastes and exposed substrata. It has exhibited Ecological Relationships and Distribution— adaptability to coarse-textured spoils, rocky substrata, Russet buffaloberry occurs as an understory in aspen, and mixed heavy-textured soils. It fixes nitrogen fir, lodgepole pine, spruce, open woodlands, and old (USDA Forest Service 1948), which explains its excel- burns (Goodrich and Neese 1982; Viereck and Little lent vigor and growth on infertile mine spoils. 1972). It often forms dense thickets on gravel bars Thornburg (1982) reported that a number of collec- bordering streams. It is distributed from east-central tions are being evaluated for use on mined lands in the Alaska to Newfoundland, and south to Maine, New northern Great Plains. York, Michigan, New Mexico, and Oregon (Viereck This species is used to restore riparian disturbances; and Little 1972). It is widespread throughout Utah at it is well suited to both mesic and dry sites. It provides elevations between 6,690 and 10,500 ft (2,000 and

642 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

apically, silvery green above, pale beneath, and clothed with trichomes that provide a silvery appearance. Flowers are yellowish and very attractive. Fruits are 0.2 to 0.31 inch (5 to 8 mm) long, drupelike, and orange to red (Welsh and others 1987). This is an attractive shrub with distinct silver-green foliage, thick leaves, and colorful fruits and flowers. Ecological Relationships and Distribution— Roundleaf buffaloberry is native to the blackbrush, ephedra, shadscale, pinyon-juniper, and ponderosa pine communities of southeastern Utah and Arizona (Welsh and others 1987). It is not reported from Colo- rado. This shrub is endemic to the Colorado Plateau (Welsh and others 1987), and occupies important areas Figure 26—Fruits of russet buffaloberry are bril- where restoration of native shrubs is desirable (fig. 27). liant red and highly attractive. Seedlings establish Investigative studies have been conducted to develop well on mine sites and other disturbances. methods to culture and establish this species by artificial plantings. Plant Culture—Seeds of roundleaf buffaloberry are relatively large compared to those of other 3,200 m) (Welsh and others 1987), and it occupies buffaloberry species. There are about 6,855 cleaned similar elevations in Colorado (Harrington 1964). seeds per lb (15,110 per kg) (Plummer and others Plant Culture—Thilenius and others (1974a) re- 1968). Abundant seedcrops are produced infrequently, ported that this shrub is well suited for dry, rocky usually once every 3 to 10 years, although some fruits banks, but experimental plantings in central Idaho on may be produced on individual plants more often. mine and roadway disturbances have not been suc- Plants flower early in spring, and fruits usually ripen cessful. Transplant stock propagated from native in early to late July (Plummer and others 1968). The stands has not established well when planted on plants are very attractive because of the color contrast adjacent disturbances. The species appears sensitive between the fruit and leaves. Berries are processed by to shade, soil disturbances, and competition from maceration to release and separate the seeds. Seeds understory herbs. It has not established well on mine germinate erratically following extended periods of spoils or on streambanks that have been subjected to wet prechilling. Young plants grow slowly, even if soil loss from flooding. Planting success may be signifi- protected from competition. Nursery and greenhouse- cantly improved with the use of better quality stock, grown stock also grow slowly, although older plants but this species has been less widely used and has not grow somewhat faster. been as successful in revegetation plantings as silver buffaloberry. Uses and Management—Russet buffaloberry occurs at rather high elevations and on sites where logging, road construction, and mining have created considerable disturbances; consequently, it has been used for revegetation of these sites. It provides useful ground cover and improves diversity in lodgepole pine forests where other understory and shrubs are lacking. Varieties and Ecotypes—None.

Family Elaeagnaceae ______Shepherdia rotundifolia Roundleaf buffaloberry Description—Roundleaf buffaloberry is a short statured plant growing 3 to 6.5 ft (0.9 to 2.0 m) in Figure 27—Roundleaf buffaloberry is a unique, height. The twigs are stellate hairy with white or attractive shrub common to arid regions of the yellowish trichomes. Leaves are ovate to oval, rounded Colorado River drainage.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 643 Chapter 23 Shrubs of Other Families

Uses and Management—This species is most im- Nevada ephedra has good tolerance to soil salinity and portant in the unique desert shrublands of southern can be found growing in well-drained to fairly poorly Utah and Arizona. It provides useful ground cover, drained soils ranging from rocky, sandy loams to fairly aesthetics, and limited, but seasonal forage for big heavy clays (Dayton 1931; Plummer 1977). Nevada game (Plummer and others 1968). Attempts to culture ephedra occurs in open stands of scattered plants, but this species have only resulted in fair success; but with commonly forms dense, slowly spreading colonies. further research, this species may be of more use for Individual plants may be connected by underground restoration of recreational sites and native shrub stolons that develop when branches are covered with communities. It has been successfully used in land- windblown sand or silt (Wallace and Romney 1972). scape plantings. It grows well with limited irrigation, Plant Culture—Seed crops are produced errati- and once established produces considerable foliage. It cally (Cronquist and others 1972; Wallace and Rom- demonstrates utility in recreation plantings for screen- ney 1972). Generally, one good seed crop is produced ing campgrounds and related sites. It also provides a every 5 to 6 years (fig. 28). Consequently, seed must be useful and attractive native shrub that can be used to harvested and stored for anticipated planting needs. restore native communities in highly used recreation Within the Great Basin, cones appear in April to early sites. May, and seeds ripen in July (Dayton 1931; Turner Some collections of roundleaf buffaloberry have not and Randall 1987). Farther south, cones appear in survived when transferred to more northern climates March and seed matures in June. Seeds are easily in central Utah. This species normally grows with a collected, cleaned, and seeded. Mature seeds must be limited understory and cannot compete well if planted collected quickly before they drop from the shrub or with a dense cover of herbs. Plants are difficult to are gathered by rodents. They are collected by beating establish from transplanting or direct seeding. the cones into a container. Establishment and mainte- Varieties and Ecotypes—None. nance of seed orchards on adapted sites might allevi- ate seed supply problems. Four to 6 years should be allowed for stand establishment before significant Family Ephedraceae ______seed crops are produced. Seeds can be easily separated Ephedra nevadensis from debris with an air screen cleaner, providing seeds Nevada ephedra with better than 99 percent purity. There are about 20,000 Nevada ephedra seeds per lb (44,100 per kg) at Description—Nevada ephedra or jointfir is a mem- 100 percent purity. ber of the gymnosperm family Ephedraceae. This Seed remains viable for long periods when stored air dioecious shrub grows to 5 ft (1.5 m) tall and has dry at room temperature. Seedlots of Nevada ephedra evergreen stems with paired scalelike leaves 0.08 to 0.2 inch (2 to 5 mm) long (Welsh and others 1987). One to several male cones are borne at the nodes. Female cones are borne solitary at nodes and produce a pair of pale-brown to yellowish-green seeds (Welsh and others 1987). The stems of Nevada ephedra are divergent, coarse, bluish-green to gray, and produced on older wood. In contrast, green ephedra produces erect, parallel, broomlike stems that are green to yellow green in color (Cronquist and others 1972; Dayton 1931). Ecological Relationships and Distribution— Nevada ephedra is a native perennial shrub that occurs throughout the Intermountain West in juniper- pinyon, big sagebrush, Indian ricegrass, rabbitbrush, black sagebrush, blackbrush, salt desert, and cresote- bush types in central and southern Utah, Nevada, Arizona, California, and Mexico (Cronquist and others 1972). It frequently grows with green ephedra; however, it is usually more abundant on drier sites and in Figure 28—Nevada ephedra generally grows on dry salt desert, black sagebrush, and blackbrush types. sites in pinyon juniper, big sagebrush, Indian ricegrass, This species is generally found growing in areas with rabbitbrush, black sagebrush, blackbrush, salt desert, 7 to 14 inches (18 to 36 cm) of annual precipitation. and creosote communities.

644 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families exhibited 89 percent germination following 15 years of Nevada ephedra has been used to stabilize rocky and storage (Stevens and others 1981a). With 25 years of sandy areas (Plummer 1977). It is well adapted for use storage, germination was 77 percent (Stevens and as a low-maintenance, low-water-requiring ornamen- Jorgensen 1994). Seeds germinate under a wide range tal for home sites, parks, recreational areas, and of temperatures without wet prechilling or other pre- administrative areas. It establishes slowly and has treatments (Young and others 1977). Seeds of Nevada not done well on exposed substrata at mine sites or on ephedra will germinate somewhat faster than seeds of road disturbances. Plants should not be grazed for two green ephedra. Nevada ephedra can be seeded sepa- to three growing seasons following seeding. Grazing rately or in conjunction with other shrubs, grasses, and trampling should be avoided until shrubs become and forbs within its native range. When drill seeded, firmly established and are 5 to 8 inches (13 to 20 cm) Nevada ephedra and most other shrubs do best if tall. planted in separate rows from competitive perennial Varieties and Ecotypes—None. grasses. Seeding Nevada ephedra through a seed dribbler or thimble seeder provides good results. Ne- vada ephedra does best when seeded on a firm seedbed Family Ephedraceae ______and covered to 0.5 inch (1.3 cm) depth. In most of the Intermountain West, Nevada ephedra Ephedra viridis should be seeded in fall to allow for effective use of Green ephedra spring soil moisture and precipitation (Plummer and others 1968). In the more southerly regions, good Description—Green ephedra is also called Brigham results have been obtained with midsummer seeding, tea, Mormon tea, or jointfir. This dioecious shrub has just prior to the summer storm period. Plants often evergreen stems, can grow to 4 ft (1.2 m) tall, and require 5 to 10 years to attain a height of more than produces a deep fibrous root system (Cronquist and 2 ft (61 cm) (Plummer and others 1968), yet seedlings others 1972; Wasser 1982). Green ephedra produces emerge and establish quickly. Seedlings withstand erect, parallel, jointed, broomlike stems that are extreme drought once established. green to yellow green (Cronquist and others 1972). Nevada ephedra can be transplanted with moderate Leaves are small ligulelike scales, and are opposite at success (Plummer and others 1977). Roots of bareroot stem nodes. Plants are dioecous; male cones are com- stock are generally weakly developed and are very pound, borne at the nodes or terminal points on the fragile and easily damaged. The root systems of con- stems. Female cones are solitary or whorled, and tainerized stock are sometimes too small to bind the sessile or peduncled (Wasser 1932; Welsh and others root plug together. The plug must be carefully re- 1987). Cones are formed in pairs and are slightly boat moved from the container to avoid breaking the roots. shaped. Low root-to-shoot ratios are common for both types of There are two varieties of green ephedra. E. v. var. stock, and survival rates are frequently less than 40 viridis has sessile female cones, nonviscid stems, and percent. Wieland and others (1971) found that Ephe- grows to 6 ft (1.8 m) tall. E. v. var. viscida has dra species could be propagated from rooted cuttings. pedunculate female cones and viscid stems. It occurs Everett and others (1978a), however, reported little on sandy soils where it may be grasslike in appear- success in rooting cuttings of either Nevada ephedra ance; it is often covered with sand with only small or green ephedra. stems protruding (Kartesz and Kartesz 1980; Welsh and others 1987). Uses and Management—Nevada ephedra pro- Ecological Relationships and Distribution— vides forage and cover for cattle, sheep, deer, and Green ephedra is a native, perennial shrub that occurs antelope (Dayton 1931; Dittberner and Olsen 1983; on shallow to medium depth soils, on sandy or rocky Kufeld and others 1973; Smith and Beale 1980; Stevens slopes and in valleys (Plummer and others 1968; 1983b; Welsh and others 1987). It is reported to be the Wasser 1982) throughout the Intermountain West most important forage plant of the ephedra species (Cronquist and others 1972). It occurs in juniper- (USDA Forest Service 1937). Use is often difficult to pinyon-sagebrush, rabbitbrush, black sagebrush, detect or measure because browsing causes the young blackbrush, salt desert shrub, ponderosa pine, and stems to break off at the joints. Nevada ephedra can mountain brush types (Plummer and others 1968; be an important source of browse during drought and Stevens 1983b) (fig. 29). Green ephedra frequently winter periods. Quail and cottontail rabbits use the grows with Nevada ephedra; however, it is usually species for food and cover (Stanton 1974). Seeds are more abundant on mesic sites with 9 to 14 inches (23 readily gathered and eaten by rodents and birds. to 36 cm) of annual precipitation. Green ephedra Rodents cache seeds, which can then emerge in clus- occurs as dense, individual bushes and in large stands. ters of new seedlings.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 645 Chapter 23 Shrubs of Other Families

seedbed. Rodents often gather planted seeds, and can limit the success of direct seedings. In the Intermountain region, green ephedra should be seeded in the fall. (Plummer and others 1968; Vories 1981). Seedlings that emerge early and quickly can best withstand extreme drought. Both green ephedra and Nevada ephedra can be transplanted with only moderate success (Luke and Monsen 1984). The roots of bareroot stock and container stock develop slowly and are easily damaged. Survival rates are frequently less than 40 percent. Uses and Management—Green ephedra provides forage and cover for cattle, sheep, deer, and antelope (Dayton 1931; Dittberner and Olsen 1983; Kufeld and others 1973; Smith and Beale 1980; Stevens 1983a; Figure 29—Green ephedra exists as an im- Welsh and others 1987). It is also grazed by bighorn portant component of many shrub communities sheep, rabbits, and quail (Stanton 1974). Green ephe- in the foothill regions of central Utah. dra is heavily hedged by sheep, particularly during winter periods. Because of its evergreen growth habit and abundant herbage that protrudes above the snow, green ephedra is an important species on winter game and livestock ranges. Seeds are gathered and eaten by It sprouts readily from the root or crown following fire rodents and birds, which tend to spread the species. (Young and Evans 1974, 1978b). Both green ephedra and Nevada ephedra can be used to stabilize rocky, sandy, and disturbed areas Plant Culture—Cones develop in April and May; (Plummer 1977; USDA Forest Service 1976; Wasser seeds ripen in July and early August. The large seeds 1982; Young and others 1979a). However, green ephe- are easily collected, cleaned, and planted. Mature dra persists much better on disturbances and exhibits seeds must be collected quickly because they are adaptation to a wider range of soil conditions. dislodged from the plant by wind. Seeds are also Although both green ephedra and Nevada ephedra quickly gathered by rodents and birds. Heavy seed have been used in ornamental and landscape plantings crops are produced erratically (Plummer and others (USDA Forest Service 1976; Wasser 1982), green ephe- 1968). Generally, one good seed crop is produced every dra has survived much better, and is less sensitive to 5 years. Consequently, seed must be harvested and supplemental irrigation and cultivation. stored for later use. Seeds are harvested by dislodging them into a container. They can be easily separated Varieties and Ecotypes—None. from debris with an air screen cleaner. Seed lots usually are 99 percent pure. There are about 25,000 Family Ericaceae ______green ephedra seeds per lb (55,100 per kg) at l00 percent purity. Arctostaphylos uva-ursi Seed remains viable for long periods when stored air Bearberry manzanita dry at room temperatures. Stevens and others (1981a) found that seedlots of green ephedra exhibited 87 Description—Bearberry manzanita is a prostrate, percent germination following 15 years of storage. mat-forming, evergreen shrub with stoloniferous stems With 25 years of storage, germination declined to 2 (Hitchcock and Cronquist 1973; Morris and others percent (Stevens and Jorgensen 1994). Seeds will 1962; Noste and Bushey 1987; Welsh and others 1987). germinate under a wide temperature range without The branches are ascending; the internodes are usu- wet prechilling or other pretreatments (Young and ally apparent, puberulent, and sometimes glandular. others 1977). The exfoliating bark has a dull brown to red color. Green ephedra can be successfully seeded with other (fig. 30). Leaves are 0.2 to 1.1 inches (0.6 to 2.7 cm) shrubs, grasses, and forbs within its native range. If long, 0.1 to 0.5 inch (3 to 12 mm) wide, oblanceolate to drill seeded, it should be planted in separate rows to spatulate, and glabrous or puberulent, especially on reduce competition with perennial grasses. Seeding the margins. The inflorescence is racemose. The axis green ephedra with a seed dribbler or thimble seeder and bracts are glandular. Flowers are pink to white, has resulted in good success. Seeds establish best and 0.16 to 0.2 inch (4 to 5.2 mm) long. Fruits are 0.2 when planted at a depth of 0.5 inch (1.3 cm) on a firm to 0.4 inch (6 to 11 mm) thick, globose, bright red, and

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periods, and thus remains as a component of various successional stages of community development. Bearberry manzanita is generally not considered fire tolerant (McKell and others 1972), but some plants are able to survive burning. Following fire, new shoots develop from portions of the plant that did not burn. Seeds stored in the soil are stimulated to germinate by the heat treatment provided from burning (Carmichael and others 1978), and numerous seedlings may appear 1 to 5 years after a burn (Pase and Pond 1964). Plant Culture—Flowers appear from March to May; fruits ripen from June to August and are dispersed from August to March (Berg 1974; Everett 1957; McMinn 1951; Munz and Keck 1959). Fruits are collected by hand stripping, but some can Figure 30—Bearberry manzanita is a mat-forming, be dislodged by beating the bush with a paddle. evergreen shrub that provides excellent ground cover. Seeds are easier to clean when the fruit is still green. If the fruit is allowed to dry, the fruit coat and fleshy material are difficult to remove. Cleaning is accomplished using a macerater to grind and remove the berrylike with separable nutlets. The four to 10 nut- fleshy material from the seeds. Grinding is fre- lets are fused or partially fused, 0.12 inch (3 mm) long, quently conducted using water to flush away the 0.08 inch (2 mm) wide, and light brown with a very fleshy material. After grinding, the material is dried thick seedcoat (Belcher 1985; Welsh and others 1987). and screened with a fanning mill to separate the seeds Many roots appear near the soil surface to form a from other materials. Certain seedlots have a large dense, fibrous system (Berndt and Gibbons 1958). number of empty seeds; these can be removed by Numerous runners extend laterally from the crown flotation or use of a gravity table separator. Seeds can to a distance in excess of 3 ft (0.9 m) (McLean 1967). be stored for long periods at room or warehouse tem- Bearberry manzanita is able to regenerate by resprout- peratures (Glazebrook 1941). Bearberry manzanita ing from stolons (Berndt and Gibbons 1958). Rowe produces between 26,800 to 58,000 clean seeds per lb (1983) reported that sprouting also occurs from large (59,080 to 127,870 per kg) (Berg 1974; Glazebrook lignotubers. 1941; McKeever 1938; Plummer and others 1968). Ecological Relationships and Distribution— Seedcoats are thick, hard, and impermeable to wa- Bearberry manzanita is circumpolar in distribution, ter; however, a small opening extends from the surface occurring in North America from Labrador to Alaska, of the seedcoat to the embryo. This is plugged with a and south to northern California, and the mountains waxlike substance that may be removed by soaking of New Mexico, Virginia, Illinois, and Nebraska (Berg the seeds in sulfuric acid. Wet prechilling treatments 1974; Sampson and Jespersen 1963). It is common in are required to overcome the embryo dormancy. In ponderosa pine and Douglas-fir forests in mountain- addition, the embryo requires afterrippening ous regions, and it also occurs in oak woodlands and (Glazebrook 1941). Most authors recommend various chaparral types in Arizona and California. Cooper and periods of acid scarification, followed by a warm pre- others (1987), Mueggler and Campbell (1986), and treatment and a period of wet prechilling. Berg (1974) Youngblood and Mauk (1985) reported bearberry and Milstein and Milstein (1976) recommended 2 to 5 manzanita was present in many of the conifer forest hours of acid scarification and planting in summer, or and aspen types in Utah and northern Idaho. In Utah a warm pretreatment at 25 ∞C (77.0 ∞F) for 60 to 120 it is encountered in conifer forests at 7,000 to 11,500 ft days, then a wet prechill for 60 to 90 days. McLean (2,140 to 3,510 m) in the north and north-central (1967) suggested an acid soak for 7 hours, followed by portions of the State (Welsh and others 1987). a warm, wet pretreatment at 68 ∞F (20.0 ∞C) for 90 Bearberry manzanita exists on a variety of soil types days, and a wet prechill at 34 ∞F (1.1 ∞C) for 90 days. including weakly developed soils and coarse-textured Giersbach (1937) recommended an acid soak for 3 to 5 materials (Carmichael and others 1978). It is usually hours, then overwinter storage outdoors in mulch regarded as a late seral species. However, it does not flats. form an abundant understory in forest communities Following treatment, seeds are usually incubated at in Utah and northern Idaho (Cooper and others 1987; high temperatures. Milstein and Milstein (1976) rec- Mueggler and Campbell 1986; Youngblood and Mauk ommended incubating pretreated seeds at a constant 1985). It persists in open and shaded areas for extended temperature of 78 to 80 ∞F (25.6 to 26.7 ∞C). Berg (1974)

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 647 Chapter 23 Shrubs of Other Families suggested either constant temperature incubation at Bearberry manzanita is widely used for landscape 77 ∞F (25.0 ∞C), or alternating 86 ∞F (30.0 ∞C) day, and plantings, and numerous selections are available. 68 ∞F (20.0 ∞C) night. These treatments produced 30 to Because it grows as a low-spreading plant, it has been 61 percent germination in 15 to 30 days. Germination used in erosion control and watershed plantings. It exceeding 90 percent has been reported by Glazebrook requires special care to assure establishment by trans- (1941). planting as container or bareroot stock. Transplants Early summer planting, following acid soaking, is must be dormant when field planted or they are recommended by Berg (1974), Milstein and Milstein susceptible to damage by frost or drought. Dormant (1976), and Swingle (1939), but early fall seeding of stock that is field planted early in the spring survives treated seeds is also employed. Seeds are small and well. should be planted approximately 0.5 inch (1.3 cm) Small rooted transplants usually grow slowly, and deep. Seed germination will normally occur in 20 to 40 often fail to survive if planted on unstable slopes or days (Milstein and Milstein 1976). On wildland sites, surfaces. Larger transplants (2-year-old stock) sur- seedling emergence has been observed to occur in 5 to vive harsh conditions much better and grow faster. 10 days. Seedlings are slow developing and can be However, growth is usually slow and plants fail to suppressed by competitive vegetation. Also, seedbeds provide immediate ground cover. Normally 3 to 5 often dry rapidly and some seedlings fail to survive. years are required before plants spread and occupy a Plants can be easily propagated by stem tip cuttings large area. If this species is used to control erosion, (Berg 1974). Doran (1957) reported that cuttings taken transplants should be spaced 1.5 to 2.5 ft (46 to 76 cm) in late fall or winter (October to February) root well. apart, depending on surface conditions. Eighty-eight percent of cuttings taken in October Collections acquired from wildland sites can be rooted in 22 weeks, but if treated with IBA, 3 mg per g propagated for field plantings, but have demonstrated of talc, 96 percent rooting occurred in 15 weeks. Stem adaptability to specific site conditions. Materials grow- cuttings taken in February rooted best, 90 percent in ing in shaded areas beneath ponderosa pine perform 11 weeks. Cuttings are rooted under greenhouse con- erratically when transplanted in open areas at similar ditions using bottom heat at 76 ∞F (24.4 ∞C). Once stem elevations. Transplants also grow poorly on road and cuttings are rooted, they survive field planting very mine disturbances, although poor performance may well. be due to poor planting stock. However, this species Bearberry manzanita is widely used for horticul- appears to be sensitive to soil conditions and has not tural plantings. Considerable information is available survived on mine disturbances. related to vegetative propagation and culture Bearberry manzanita slowly invades road and log- (Darbyshire 1971; Dehgan 1972; Dehgan and others ging disturbances, but only in areas where some top- 1975; Hildreth 1969; Leopold and Kriedeman 1975). soil has accumulated. Although use is questionable on Dehgan and others (1977) recommended stem cut- harsh disturbances, its growth habit is an important tings be taken in winter or early spring and treated feature. Topsoiling and use of soil amendments sig- with low IBA concentrations. Rooted cuttings survive nificantly enhance survival and growth rates. better if hardened off gradually, starting with com- This plant has adapted well when used for recre- plete shade and progressing to an open situation. ational sites, landscaping summer homes, and other Potting material and pots should be well drained to low-maintenance landscape areas. Considerable dif- avoid overwatering. ferences in ecotypes and growth forms are available to provide diversity in landscape plantings. It could be Uses and Management—Various authors con- more widely used within the Intermountain area for sider the genus, Arctostaphylos, to provide quality this purpose. browse (Conrad 1987), but reports by Cada (1971), Cooperrider (1969), Cowan (1947), Gullion (1964), Hill Varieties and Ecotypes—Various horticultural and Harris (1943), Kamps (1969), Kufeld and others selections are available, but materials have not been (1973), and Lovaas (1958) indicated that the plant selected for wildland uses. receives moderate use by wildlife. In general, it is one of the earliest shrubs to initiate growth in spring. Leaves are also maintained on the shrub during win- Family Fagacae ______ter, and game animals are attracted to the plant Quercus gambelii during the winter and spring periods. It often grows on Gambel oak open sites, not heavily covered by snow, that are important winter grazing locations. Game animals Description—Gambel oak (fig. 31) is a member of frequently utilize the shrub, although it may not be a the white oak subgenus Lepidobalanus. Most taxono- major portion of their diet. mists consider Gambel oak part of the Quercus undulata complex, which hybridize readily (Welsh

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1986). Gambel oak is a native, broadleaf, deciduous Wasatch Mountains, Wasatch Plateau, and west slope shrub that typically forms dense stands (Neilson and of the Rocky Mountians in western Colorado in nearly Wullstein 1983). It generally grows from 3 to 20 ft (0.9 continuous belts (oakbelt) between 5,500 and 7,000 ft to 6 m) in height, forming spreading thickets con- (1,700 and 2,100 m) (Cottam and others 1959). Gambel nected by underground rhizomes (Christensen 1949). oak generally requires annual precipitation between In the southern end of its range it can be treelike and 15 and 22 inches (38.1 and 55.9 cm) (Harper and as much as 20 to 25 ft (6 to 7.6 m) tall (Wagstaff 1985). others 1985). However, it does occur in some areas Leaves are alternate and broadly obovate to oblong where annual precipitation is as low as 12 inches lanceolate, with considerable variation among clones. (30.5 cm). The most common soils in which Gambel Leaves are somewhat leathery, yellow green to dark oak occurs are calcarcous, heavy, and fine-grained green above and pale yellowish and densely hairy loams. It can also be found on sandy, gravelly, and clay below. They turn a deep-reddish color in fall. loams, and alluvial sands (Christensen 1955; Tucker This monoecious, wind-pollinated shrub bears red and Muller 1958). pistillate flowers in the upper leaf axils. Male flowers Gambel oak occurs as a climax species on many sites, grow in pendent catkins that are borne singly or in and as a mid or early seral species on others (Harper groups at the base of new shoots (Harper and others and others 1985). In some areas Gambel oak is consid- 1985). Fruits are brown acorns enclosed in a cap. ered the ecological equivalent of ponderosa pine (Harper Acorns form and mature in 1 year (Harper and others and others 1985). 1985). Reproduction is primarily vegetative from an Plant Culture—Gambel oak reproduces both veg- extensive, freely branching, underground network of etatively and sexually. Clones expand through both lignotubers (enlarged, stemlike structures covered by radial and lateral growth (Christensen 1955). Rate of numerous adventitious buds), roots, and rhizomes vegetative spread ranges from 1.5 to 12 inches (381 to (Tiedemann and others 1987). Lignotubers account for 360 mm) annually; the average is 4 inches (10 cm) per up to 72 percent of the total belowground biomass year (Christensen 1955). Gambel oak sprouts quickly (Tiedemann and others 1987). after burning, wood harvesting, chaining, cabling, or Ecological Relationships and Distribution— other disturbances. Sprouts may appear 10 days after Gambel oak is widely distributed throughout the South- removal of top growth (Harrington 1985). Rate and ern and Central Rockies. In the Intermountain West, degree of sprouting are greatest from clones with Gambel oak occurs in valleys, foothills, on alluvial smaller stems. Clones with stems greater than 12 fans, and on lower mountain slopes between sage- inches (31 cm) in diameter are poor sprouters (Reynolds brush, pinyon-juniper, aspen, and spruce-fir types. and others 1970). When it is not the dominant species, it can be found Sexual reproduction is by acorns. Acorns develop growing as a codominant with big sagebrush, pinyon, and mature in 1 year (Harper and others 1985). They juniper, ponderosa pine, mountain mahogany, bitter- usually ripen in late September and October, and can brush, cliffrose, aspen, snowberry, spruce, and fir. be collected by beating them into a container. Acorns Gambel oak occurs between 4,000 and 9,000 ft must be stored in a cool, moist environment or they (1,200 and 2,700 m). Pure stands occur along the will dry and die. Storage should not extend beyond the first winter following collection because of rapid loss of viability. Seeding should occur in late fall or early spring. High pregermination mortality can result from predation by birds, mammals, insects, and parasitism. Acorns can germinate in fall after dispersal; they may be killed by frost if not covered by litter and snow (Christensen 1949). Seedlings are susceptible to damage by early fall or late spring frosts, grazing, and summer drought (Neilson and Wullstein 1983). Gambel oak seedlings are rare in many areas, particularly in the northern part of its range (Christensen 1955; Reynolds and others 1970). Plants can be established from properly planted container stock. Establishment success has also resulted from moving partial clones with front-end loaders. This involves moving and placing 2 to 3 ft Figure 31—Clumps of Gambel oak provide soil (61 to 90 cm) of soil, roots, rhizomes, and lignotubers. protection and habitat for wildlife and livestock. Gambel oak wildings do not transplant well.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 649 Chapter 23 Shrubs of Other Families

Uses and Management—Gambel oak and associ- band tailed pigeons, squirrels, and numerous small ated species provide valuable food and cover for live- mammals and birds (Christensen 1949; Harper and stock and many wildlife species. This species is gener- others 1985; Reynolds and others 1970; Steinhoff ally not highly palatable, but its abundance and 1980). Many small birds and mammals cache acorns availability, particularly on fall, winter, and spring for later use. ranges, make it an important browse species. Gambel Gambel oak is a valuable browse and cover plant in oak is regarded as fair forage for all classes of livestock many areas, but it can grow too dense and exclude (Harrington 1985). Summer cattle use has been iden- understory vegetation, big game, and livestock. A tified in northern Utah (Julander and Robinette 1950). mosaic of clones or stands interspersed with open Sheep use can be light (Dayton 1931). Gambel oak is areas provides the best habitat for big game and used extensively by mule deer throughout the year livestock (Stevens and Davis 1985). Management ob- (Bowns 1985; Harper and others 1985; Kufeld and jectives should be directed toward maintaining a mo- others 1973). Elk browse oak during winter and spring saic of clones and interspaces. Fire, herbicides, and (Kufeld 1973; Reynolds and others 1970). Bighorn mechanical treatment can be used successfully to open sheep make use of the plant during summer months stands and increase accessibility to clones (Bowns (Rominger and others 1988). 1985; Stevens and Davis 1985). When Gambel oak is Young shoots and new growth are the most pre- disturbed, understory species should be seeded to ferred portions of this shrub. Preference of most brows- improve composition and production and to retard oak ing animals for Gambel oak varies with availability of suckering and regrowth (Stevens and Davis 1985). other species. Use increases as other species become Wood value of Gambel oak can be high (Clary and less available. Gambel oak can be poisonous to live- Tiedemann 1986). Average energy content is esti- stock when used exclusively, but when consumed in mated at 340,000 Btu per ft2 (4.77 kcal per g) (Harper combinations with other species, it is generally harm- and others 1985); it can be an economically important less (James and others 1980). fuel wood with high heat-yielding qualities. Gambel Quantity and quality of associated forage produc- oak has been used successfully for long-term rehabili- tion can be very high in Gambel oak communities. tation of disturbed sites (Dittberner and Olsen 1983). Understory production may reach 3,000 lb per acre Gambel oak resprouts rapidly following fire. Rate of (3,360 kg per ha). Understory production within oak recovery varies with fire severity, climatic factors, and clones can be twice as high as between clones. Under- site characteristics (Harper and others 1985). Fire story species within clones also green up 2 to 3 weeks generally does not harm or kill Gambel oak. Spring earlier in the spring and stay green later in the season and fall burns have little detrimental effect. However, than do the same species between clones (Bowns 1985; summer burns can inhibit regrowth and may even kill Stevens and Davis 1985). Gambel oak (Harrington 1985). Following fire, the In most areas the Gambel oak type has the highest rate and amount of resprouting can be reduced by forage productivity potential of any vegetative type in seeding competitive herbs (Stevens and Davis 1985). the Intermountain West. It has been severely abused Gambel oak generally sprouts as early as 10 days after by livestock at many locations. Productive understory being burned (Harrington 1987). When Gambel oak is species have been grazed out and replaced by less burned, chained, cabled, or otherwise disturbed, deer productive plants. In most areas the understory is and elk browsing increase (Stevens and Davis 1985). almost nonexistent. As a result oak density and clone Deer, elk, and livestock seem to prefer burned over oak size have increased substantially. communities (Harper and others 1983). The Gambel oak type is important for livestock, Improved Varieties—None. Considerable varia- wildlife, and watershed protection. These areas can be tion exists with this species throughout its geographical critical spring and fall ranges for cattle, sheep, deer, range. and elk, and winter range for elk. Floods produced by high-intensity July and August rainstorms are common on depleted Gambel oak stands. Heavy snow pack Family Oleaceae ______and spring snowmelt have the potential for producing destructive spring runoff from these depleated com- Fraxinus anomala munities. It is important that the Gambel oak type be Singleleaf ash well managed and restored to its productive potential. Gambel oak also provides valuable escape, thermal, Description—Singleleaf ash is one of only a few and travel cover for deer, elk, livestock, and small native ashes to occur in the Intermountain region. It is mammals. A large number of birds use Gambel oak for a small, deciduous shrub or tree, commonly 8 to 13 ft nesting, hiding, and resting (Marti 1977). Acorns are (2.4 to 4 m) tall (Welsh and others 1987), but some- highly preferred by deer, elk, hogs, javelina, turkey, times attaining heights of 26 ft (8 m) (Goodrich and Neese 1986; Harrington 1964). It usually develops

650 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families multiple stems, and branchlets that are characteristi- Plant Culture—Small, rather inconspicuous flow- cally four angled. Leaves are simple to two or three ers appear from April to May at about the same time foliate, ovate, crenate-serrate to subentire, glabrous, the leaves develop. Flowers are grouped in terminal or 0.6 to 2.6 inches (1.5 to 6.5 cm) long, and 0.4 to 2.4 axillary clusters. Fruits are long, single-seeded sama- inches (1 to 6 cm) wide. Flowers are polygamous and ras with distinct wings that are borne in clusters. occur in panicles with a minute calyx and no corolla. They ripen from mid-July to mid-August (Plummer Fruit is a one-seeded samara, 0.5 to 1 inch (12 to 25 and others 1968). As fruits mature they change from mm) long, 0.31 to 0.4 inch (8 to 10 mm) wide, and green to yellow or light brown. Fruits often remain on winged almost to the base (Goodrich and Neese 1986; the bush for a number of weeks before dispersing. Harrington 1964; Kearney and Peebles 1942; Munz Ripened fruits can easily be collected by hand picking and Keck 1959; Welsh and others 1987). or dislodging them onto canvas spread on the ground. Fruits are dried before cleaning. They are separated Ecological Relationships and Distribution— from stems and leaves by fanning or with air screen Singleleaf ash occurs from Colorado west to California cleaners. The wings can be removed from the fruit by and south to New Mexico and Arizona (Harrington grinding or maceration. Removing the wings aids in 1964). It is common in west-central and southwestern planting, but does not affect germination. Seeds of Colorado at 4,500 to 6,000 ft (1,400 to 1,800 m) most ash species must be stored in sealed containers at (Harrington 1964). It occurs on rocky, sandy canyons 4l F (5.0 C) and 7 to 10 percent moisture content. along the Green River in Utah (Goodrich and Neese ∞ ∞ Singleleaf ash produces good seed crops infrequently. 1986), but is more widespread throughout eastern and Abundant crops may be produced once every 3 to 5 southern Utah. It exists in pinyon-juniper woodlands years. Insects damage a number of seeds as they ripen. and mixed desert shrub communities mainly on rim- Usually, less than 50 percent of the seeds from wild- rock or along drainages at elevations between 2,900 land collections are viable. A number of freshly col- and 8,500 ft (890 and 2,600 m) (Welsh and others lected seeds will germinate, but most are dormant and 1987). It occupies dry canyons and gulches between require wet prechilling. Seeds sown in fall usually 3,000 and 11,000 ft (910 and 3,400 m) in California receive an adequate wet prechilling treatment and (Munz and Keck 1959). germinate uniformly in spring. Artificial wet prechill- The plant is important on upland sites where it ing of ash seeds is accomplished by diurnal alternating grows intermixed with salt desert shrubs, pinyon- temperatures of 86 and 68 F (30 and 20 C) (Bonner juniper, and basin big sagebrush. It is particularly ∞ ∞ 1974b). abundant in the Four Corners region and throughout Uncleaned or dewinged fruits are difficult to seed much of the Colorado River drainage where it occurs with conventional drills. The fruits are large and on sandstone outcrops and shallow soils (fig. 32). It irregular in shape and do not flow through seedboxes does not form dense stands, but is normally found and seeding equipment. Fruits should be planted in scattered among rocky bluffs and ledges. It also occurs the soil at a depth of about 0.5 inch (1.3 cm). on limestone soils, but usually on well-drained sites Trial seedings, primarily in central Utah, have pro- (Stark 1966). duced irregular stands. This has been attributed to seeding poor quality seed. Seedlings and young plants grow quite slowly. Even under irrigation, nursery- grown plants develop slowly. Wildland planted seedlings often increase only 6 inches (15.2 cm) in height during a single season. As plants reach 3 to 5 years of age, the growth rate increases. A dense, well-branched root system is formed the first year; once established the plants are very persistent. Singleleaf ash can be grown as transplant stock for field plantings, although transplanting success has been quite erratic. One-year-old transplants are not very large, usually less than 5 to 6 inches (12.7 to 15.2 cm) tall, but with healthy root systems. Survival of transplanted stock has often been 25 to 40 percent, but losses are often attributed to extreme conditions of the planting sites. Growth response of singleleaf ash has varied among collections. Plant materials ob- Figure 32—Singleleaf ash occurs on shallow soils and tained from eastern Utah have grown quite slowly, but sandstone outcrops in the Four Corners area. plants establish and persist very well.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 651 Chapter 23 Shrubs of Other Families

Uses and Management—Singleleaf ash is one of shrubby species generally produce semierect stems. only a few shrubs or trees adapted to rocky and arid Flowering stocks are herbaceous, erect, and leafless. sites in the sagebrush, salt desert shrub, and pinyon- Flowers are small and arranged in umbels, cymes, or juniper communities. It develops an upright growth racemes. The fruit is a three-angled or three-winged form that provides food and cover for wild animals and achene (Dayton 1931, 1960; Hitchcock and others domestic stock. The plants are very persistent, and are 1964; Welsh and others 1987). Sulfur eriogonum, not eliminated by extended periods of drought or Wyeth eriogonum, and Wright erigonum have been heavy browsing. Singleleaf ash is also well adapted to the most successful species used in restoration projects. infertile soils, and performs well in such situations. They are easily cultured and establish well without The plant serves as an overstory species with sage- extensive seedbed preparation or protection of young brush and fourwing saltbush, providing cover and seedlings. They exist on dry situations, and prefer protection for wildlife in barren areas. When inter- rocky, sandy, well-drained soils in areas of moderate mixed with pinyon and juniper, it provides forage to low rainfall. Consequently, they are useful shrubs diversity for game and livestock. in restoration of harsh sites. Singleleaf ash competes well with pinyon and juni- Varieties and Ecotypes—None. per, and if not heavily grazed, it is able to remain vigorous and healthy amid dense stands of these trees. The species is an important plant for watershed pro- Family Polygconaceae ______tection; it grows in areas where high-intensity rains occur infrequently, but can cause considerable ero- Eriogonum heracleoides sion. This species is also able to persist and provide Wyeth or whorled eriogonum, buckwheat cover on sites where few other species may occur. Singleleaf ash has been used to a limited extent on Description—Wyeth eriogonum is a native peren- mine sites, roadways, and recreational disturbances. nial, mat-forming subshrub with woody branches and A slow growth rate, however, limits its value in areas persistent vegetative stems. Stems arise from where rapid development of cover is desired. It is rosettelike bases and produce distinctive whorled suited to rocky soils, but is not well adapted to mine leaves. Flowering stems are densely branched and disturbances if planted on mine wastes or substrates. grow to 2.5 ft (76 cm) in height. Cream-colored to yellow It has considerable potential for landscape plantings, flowers develop on umbellate inflorescences (fig. 33) particularly in highly scenic areas of eastern and (Hitchcock and others 1964; Welsh and others 1987). southern Utah. It can be used to restore disturbances Abundant seeds are generally produced each year. near recreational areas, parks, and roadways. Addi- There are about 141,000 seeds per lb (310,800 per kg) tional information is needed to improve the culture of at 100 percent purity. this important species. Ecological Relationships and Distribution— Wyeth eriogonum is widely distributed in British Varieties and Ecotypes—None. Columbia, Montana, Utah, Wyoming, Nevada, and California (Dayton 1960). It is most prevalent in moun- Family Polygconaceae ______tain brush and pinyon-juniper communities, but it also occurs in sagebrush-grass, aspen, and spruce-fir Eriogonum heracleoides, Wyeth or types. It invades disturbed sites but is less competitive whorled eriogonum, buckwheat with annual weeds, particularly cheatgrass, than sul- Eriogonum umbellatum, Sulfur fur buckwheat. This species does well on disturbed eriogonum, buckwheat sites and openings in various communities and will Eriogonum wrightii, Wright eriogonum, spread naturally if not heavily used (Plummer 1970). It does increase following burns. buckwheat Plant Culture—Wyeth eriogonum, sulfur erigonum, Introduction—Eriogonum or buckwheat, as most and Wright eriogonum produce good seed crops most species are commonly called, occurs only in North years. Even during dry years, some seeds develop. America. There are a considerable number of Seeds are quite easily collected by beating them into a Eriogonum species in the Western United States (Day- container. Seeds are not difficult to clean; they are ton 1931, 1960; Harrington 1964; Hitchcock and oth- simply separated from the debris with a fan and screen ers 1964; Judd 1962; Welsh and others 1987). The separator. genus includes annual and perennial herbs, sub- Seeds of all three species can be drilled or broadcast shrubs, and woody shrubs. Most species have tap- seeded, but they should not be planted more than 0.1 roots; some of the subshrubs have spreading or pros- inch (3 mm) deep. Fall and winter seeding are pre- trate stems that root at the joints (Dayton 1960). The ferred. Seeds of all three species germinate readily,

652 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

reduced. Birds and small mammals seek out and consume the seeds. This species has considerable ornamental potential, especially on low-maintenance areas. Improved Varieties—None.

Family Polygonaceae ______Eriogonum umbellatum Sulfur eriogonum Description—Sulfur eriogonum or buckwheat is a perennial subshrub with a strong taproot and freely branching crown. The branches can be prostrate and mat forming, but are usually upright, growing to a height of 1 ft (30.5 cm). Leaves are oval to oblanceolate, green above and grayish beneath, arising from rosettelike stem tips. The inflorescence is a freely branching umbel on a stem about 1 ft (30.5 cm) long. Leaflike bracts develop near the middle of the flowering stem. Flowers are cream colored to deep yellow or greenish yellow. Flowering stems and floral tissue often persist throughout the winter (Dayton 1931, 1960; Hitchcock and others 1964; Welsh and others 1987). Ecological Relationships and Distribution— Sulfur eriogonum occurs widely throughout Western North America in big sagebrush, pinyon-juniper, mountain brush, ponderosa pine, spruce-fir, aspen, and subalpine communities (Dayton 1960). It is especially Figure 33—Wyeth eriogonum flowers in early sum- well adapted to rocky and exposed sites, and quickly mer and generally produces a reliable seed crop. invades and occupies disturbed areas (fig. 34). Seedlings are persistent and competitive. They grow rapidly and

resulting in fairly uniform emergence over a 2-week period in early spring. Seedlings are fairly frost and drought tolerant. All three species establish fairly well when planted in combination with grasses, shrubs, and forbs. Aggressive perennial grasses can, however, become dominant. Sulfur, Wyeth, and Wright eriogonum establish well from direct seeding and spread naturally on severely disturbed sites such as mines, road cuts and fills, construction sites, and gravel and sand pits. These species are generally able to persist and increase in density following wildfires. Buckwheats most often increase in importance. Uses and Management—Livestock and big game graze Wyeth eriogonum (Dayton 1960). Most use occurs in late fall and winter when other species are dormant, but some use occurs during summer. This species recovers well from heavy use and will increase Figure 34—Sulfur eriogonum directly seeded on in density and productivity when grazing pressure is shallow soils persists with competitive annual grasses.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 653 Chapter 23 Shrubs of Other Families will spread into cheatgrass and perennial bunchgrass burroweed and snakeweed (Judd 1962). Birds and communities. small mammals seek out and consume the seeds. Wright eriogonum persists well during drought peri- Plant Culture—Seeds of sulfur eriogonum can be ods and will increase following burns. This species has harvested, processed, and seeded as described for been seeded successfully on disturbed sites. It can Wyeth eriogonum. provide rapid soil stabilization and will spread fairly Uses and Management—Sulfur buckwheat can aggressively from seed. be an important browse species for sheep, goats, cattle, mule deer, elk, and antelope (Dayton 1931, 1960; Varieties and Ecotypes—None. Kufeld 1973; Kufeld and others 1973; Monsen 1975; Smith and Beale 1980). This shrub is semievergreen Family Ranunculaceae ______and produces useful forage. Plants receive some summer use, but most use occurs in late fall and winter. Clematis ligusticifolia Even when heavily grazed, sulfur eriogonum survives Western virginsbower and spreads on sites where other important forage plants have succumbed (Monson 1975). Birds and Description—Western virginsbower is a woody, small mammals consume the seeds. climbing vine. The vigorous, glabrous to densely stri- The species has been seeded successfully on severely gose, or villous stems grow to 33 ft (10 m) in length, disturbed sites. It provides rapid soil stabilization and forming masses of vegetative growth that clamber will spread fairly aggressively from seeds to fill open- over neighboring trees and shrubs. Root systems are ings in the community. It is able to survive on unstable shallow and fibrous. The opposite deciduous leaves soils, but is not able to persist if buried by soil slough- are pinnately compound with 3, 5, or 7 lanceolate to ing. Like other species of eriogonum, sulfur eriogonum broadly ovate to chordate leaflets. Each leaflet is 0.8 to has considerable ornamental value, especially in low- 3.0 inches (2 to 7.6 cm) long and coarsely few toothed, maintenance, low-precipitation situations. or sometimes lobed or nearly entire. The twisting petioles develop from enlarged stem nodes and curl around Varieties and Ecotypes—“Sierra” sulfur flower supporting vegetation (USDA Forest Service 1937). buckwheat (E. umbellatum var. polyanthum) was re- The sweet-smelling dioecious flowers are clustered in leased for use in California. few-to-many-flowered bracteate cymes. The four oblong- lanceolate sepals are white to cream colored, 0.2 to Family Polygconaceae ______0.6 inch (5 to 15 mm) long. There are no petals. Male flowers have numerous stamens and no pistils. Sta- Eriogonum wrightii mens of the female flowers are numerous, but sterile. Wright eriogonum Pistils are numerous in a capitate cluster. The plumose styles of the villous achenes elongate to 1.0 to 2 Description—Wright eriogonum is a low, white, inches (2.5 to 5.1 cm) at maturity (Clebsch 1979; woolly, perennial shrub usually less than 2 ft (61.0 cm) Harrington 1964; Hitchcock and others 1961; Welsh tall. It commonly forms a basal crown from which and others 1987). Chromosome number is 2n = 16. many branched stems emerge. The leaves are oblan- Geographic races are sometimes recognized (Vines ceolate, sharp pointed at the tips, and covered with 1960). fine, white, woolly hairs. The white or pinkish flowers Ecological Relationships and Distribution— are borne in umbels at the apex of rather short leafless There are more than 200 species of Clematis, occur- flower stalks (Judd 1962; Welsh and others 1987). ring primarily in the Temperate Zone of the Northern Ecological Relationships and Distribution— Hemisphere (Hitchcock and others 1964; Rudolf 1974). Wright eriogonum is found primarily in pinyon-juniper, Western virginsbower is distributed from British Co- oak woodlands, and chaparral types in southern Utah, lumbia west of the Cascades to the Columbia River, California, Arizona, New Mexico, and Texas (Welsh south on both sides of the Cascades to southern Cali- and others 1987). fornia, and east from Saskatchewan to New Mexico Plant Culture—This species is easily established (Hitchcock and others 1961; Rudolf 1974; Welsh and by direct seeding. Seedlings are vigorous, drought others 1987). It commonly occurs along drainages, in tolerant, and persistent. Seeds are usually hand har- riparian communities, and on steep talus slopes and vested and processed with an air separator. canyon walls where it trails over boulders, banks, and vegetation. Uses and Management—Wright eriogonum is con- Western virginsbower is common on roadsides and sidered fairly good forage for sheep and goats and fair other disturbed sites. It frequently occurs with other for cattle (Judd 1962; USDA Forest Service 1937). woody species such as cottonwood, willow, common Under heavy grazing, this species can be replaced by

654 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families chokecherry, Rocky Mountain maple, bigtooth maple, were quite sensitive to competition from cheatgrass rubber rabbitbrush, ponderosa pine, and Douglas-fir. and broadleaf herbs. It grows on well-drained sandy to rocky soils, ranging Bareroot and container stock establish readily and from weakly acidic to moderately basic, and may grow rapidly. Seeds should be cleaned carefully to occur on somewhat saline soils (Wasser 1982). It improve stand uniformity in nursery plantings. Con- usually grows in areas receiving 12 to 20 inches (30.5 tainer seedlings may be established from seeds or from to 50.8 cm) of annual precipitation, but it is most cuttings of partially matured wood harvested from abundant in areas receiving supplementary water. late spring to late summer. Young wood with short Drought tolerance of established plants varies among internodes gives less satisfactory results. Leaf bud populations. Most are weakly to moderately drought cuttings taken in summer may be rooted under mist tolerant (Wasser 1982). Seeds and vegetative produc- (Hartmann and others 1990). Layering is useful if only tion are markedly reduced in dry years. a few plants are needed. Plant Culture—Flowering occurs from March to Uses and Management—Western virginsbower August and achenes ripen from May to December, is used in seeding or transplanting projects to im- depending on geographic location (Mirov and Kraebel prove cover and forage for many birds and small 1939; Plummer and others 1968; Swingle 1939). Mod- mammals. Smith (1953) found that captive mule deer erate to heavy seed crops are produced almost every preference for western virginsbower was greatest in year. Mature fruits are dry and brown with feathery early summer and decreased considerably by late white styles. They are dispersed by wind and gravity; summer relative to other native shrub forages pro- rate of dispersal depends on weather conditions. Fruits vided. The plant receives little use by big game on are gathered by hand or with a vacuum seed harvester mule deer winter ranges in southern Idaho (Holmgren (Plummer and others 1968). Dried fruits are hammer- 1954). Dixon (1934) reported that its palatability to 3 milled with a ⁄16-inch (0.5-cm) screen operated at mule deer in California was low. It receives little use 1,120 rpm to detach the styles from the achenes by livestock. (Wasser 1982). Debris is removed by fanning. Seeds Because of its rapid growth rate, generally low are not removed from the achenes. palatability, and ability to naturally invade disturbed Estimates of the number of seeds per lb range from sites, western virginsbower is a potentially valuable 93,000 to 328,000 (205,000 to 723,000 per kg) (Mirov species for restoring disturbed sites with native veg- and Kraebel 1939; Rudolf 1974; Swingle 1939). A etation (Bailey 1947; Plummer and others 1968; Rehder purity of 20 to 50 percent and germination of 70 1940; Van Dersal 1938). It has been used for soil percent are recommended for seed purchases. Greater stabilization on riparian areas and mined lands purities are difficult and expensive to achieve. Viabil- (Thornburg 1982), and is particularly useful on steep ity has been maintained for 2 years in dry storage slopes that would otherwise be revegetated only slowly (Plummer and others 1968). by herbaceous species (fig. 35). Wider use of the species Wet prechilling at 33 to 40 ∞F (0.6 to 4.4 ∞C) for 2 to 180 days is required to release embryo dormancy (Heit 1968a; Rudolf 1974; Young and Young 1986). Germination of wet prechilled seeds after 40 to 60 days at 68 to 86 ∞F (20 to 30 ∞C) ranged from 11 to 84 percent (Mirov and Kraebel 1939; Plummer and others 1968; Swingle 1939). Western virginsbower may be direct seeded in mixtures with other shrubs by drilling or broadcasting. Untreated seeds may be fall planted (Rudolf 1974). Seed should be wet prechilled prior to spring planting (Swingle 1939). Seeds should be covered with about 0.3 to 0.5 inch (7 to 13 mm) of soil. Seedlings are vigorous, but growth is reduced by herbaceous competition. Consequently, no more than 50 percent of the mixture should consist of grass seeds. Western virginsbower normally should be seeded in mixtures at a rate of 1 lb per acre (1.1 kg per ha) or less (Wasser 1982). Holmgren (1954) reported good growth rates, but low survival of seedlings emerging from seedings Figure 35—Western virginsbower growing on a on southern Idaho big game winter range. Seedlings road fill provides excellent soil stabilization.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 655 Chapter 23 Shrubs of Other Families is precluded by the difficulty of harvesting and pro- In general, species of Ceanothus have a number of cessing large quantities of seed. Seedlings established attributes that are important for revegetation. Seeds from seeds or transplant stock develop rapidly, often are easily cleaned and planted. Small seedlings are flowering the first year, and provide cover while other vigorous and young plants grow rapidly; only a few planted species develop more slowly. Seeds and seed- other shrubs grow as quickly as Ceanothus. Species of lings are sometimes consumed or damaged by small Ceanothus are capable of nitrogen fixation and most mammals. Leaf spots and rusts are often noted on the are well adapted to infertile soils. Most are excellent foliage, and downy mildew is occasionally a problem forage plants, yet recover well from grazing. (Holmgren 1954). Plants are moderately shade and frost tolerant, but thermal tolerance varies among ecotypes (Wasser Family Rhamnaceae ______1982). Shoots may be destroyed by wildfires. Some Ceanothus cuneatus plants survive by resprouting from rootstocks. The Wedgeleaf ceanothus species also reestablishes burned areas by wind dispersal of seeds produced offsite (Plummer and others Description—Wedgeleaf ceanothus (fig. 36) is an 1968; Thornburg 1982). evergreen shrub (Munz 1974) distributed from Mexico Western virginsbowers’ cascade of yellowish foliage to California, Nevada, Oregon, and southern Wash- and clusters of feathery styles make it a unique and ington (Munz 1974; Peck 1941). It is a highly palatable attractive ornamental for use in dry areas (Plummer evergreen species, and is heavily browsed as a winter and others 1968). The plant requires a large shrub or forage (Conard and others 1985; Gibbens and Pieper small tree for support. It may be trained to grow over 1962; Gibbens and Schultz 1963). It is an upright a fence or trellis. Late winter pruning confines the shrub ranging in height from 3 to 23 ft (0.9 to 7 m) plant to smaller areas (Clebsch 1979). Plants are most (Munz 1974) and forming dense thickets (Stubbendieck productive if planted well away from the supporting and others 1986). plant in slightly acid to moderately alkaline soil. Rich, moist soil and mulching provide needed thermal pro- Ecological Relationships and Distribution— tection for the root system. Collections from Butte, Shasta, and Siskiyou Counties in California, and Jackson County, OR, have been Varieties and Ecotypes—None. Selection trials planted throughout the Intermountain area. Selec- are being conducted by the USDA Soil Conservation tions from approximately 2,000 ft (610 m) elevation, Service, Pullman Plant Materials Center, Pullman, growing on granitic and basalt soils, have been planted WA, for a cultivar adapted to the northern Intermoun- in Utah, Idaho, Nevada, Wyoming, and Colorado. This tain region. species occurs as an understory in a number of ponderosa pine communities; outplantings have primarily Family Rhamnaceae ______been confined to ponderosa pine or mountain brush sites in the Intermountain area. Various collections Ceanothus species have also been used to treat disturbed sites in southern California (Fessenden 1979). Introduction—Numerous species of Ceanothus occur in the Western United States. Munz (1974) described over 75 species in California. Considerably fewer species occur in the Intermountain region (Harrington 1964; Welsh and others 1987). However, many species common to the Southwest, northern California, Oregon, and southern Washington have been tested in the Intermountain area. Species that have been entered into adaptation trials were selected from sites with somewhat similar climatic and edaphic conditions. Different species of Ceanothus possess important traits and have been planted in an attempt to enhance wildlife, range, watershed, or horticultural resources in areas outside their natural range. Al- though a number of species have been initially screened for adaptability to the Intermountain re- Figure 36—Capsules of wedgeleaf ceanothus must gion, the species discussed in this chapter have been be harvested when green as they dehisce explosively the most promising. when dry.

656 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Plant Culture—Seeds of the approximately 12 Uses and Management—The species demonstrates collections tested in the Intermountain area express usefulness for seeding disturbed watersheds; it grows similar features. They are small with 49,000 to 54,000 rapidly and furnishes excellent ground cover. It also cleaned seeds per lb (108,000 to 119,000 per kg) (Reed provides cover on open, wind-blown game ranges. This 1974; Van Dersal 1938). Plants established in the species’ ability to fix nitrogen also contributes to its Intermountain area usually produce good seed crops usefulness in range, watershed, and disturbed land annually from healthy bushes. Few seedlings have plantings. appeared from natural spread in any study site, al- Varieties and Ecotypes—None. though some sites are frequently cultivated, which would result in seed burial. Seeds germinate and establish well from late fall Family Rhamnaceae ______and early winter seedings. Rodents gather ripened seeds from trial plantings and native stands (Conard Ceanothus greggii and others 1985). Consequently, late fall plantings are Desert ceanothus advised to reduce rodent damage to new plantings. Wedgeleaf ceanothus seeds germinate abundantly Description—Desert ceanothus is a low-growing after fires (Reed 1974; Sampson and Jespersen 1963); to erect shrub 1 to 6.5 ft (0.3 to 2 m) tall, with thick, seeds also benefit after being treated with boiling persistent, entire leaves (Welsh and others 1987). water (Grisez and Hardin 1967). Seeds placed in Desert ceanothus grows in mixed desert shrub com- boiling water for 1 to 3 minutes germinated rapidly munities including pinyon-juniper, ponderosa pine, and more uniformly than untreated seeds. This re- chaparral, and mountain brush sites in Nevada, Cali- sponse has been recorded for numerous collections. fornia, Arizona, New Mexico, Texas, and Mexico Seeds that are fall seeded following hot water treat- (Kearney and Peebles 1960; Thornburg 1982; Welsh ment are generally adequately wet prechilled and and others 1987). It occurs at elevations between 4,000 germinate well in the spring. and 9,400 ft (1,220 to 2,870 m) in southern Utah New seedlings are quite vigorous and grow rapidly. (Welsh and others 1987). Growth exceeds that of big sagebrush, antelope bitter- Ecological Relationships and Distribution— brush, and other commonly seeded shrubs. Young Only a limited number of collections, primarily from plants are able to recover following some browsing, southern Utah, have been used in wildland seedings. but Medin and Ferguson (1980) reported that develop- Collections acquired from scattered stands growing on ing stands established by seeding and transplanting rocky slopes, and from nearby pure stands, have been on winter game ranges in Idaho ultimately succumb to tested. Although this species grows on a variety of soil continued heavy grazing by big game, small mam- types (Carmichael and others 1978; Plummer 1977; mals, grasshoppers, and to winter injury. Plants are Stark 1966; Thornburg 1982), collections obtained very palatable (Gibbens and Schultz 1963; Scrivner from basic to coarse-textured soils have been the only and others 1988) and are heavily used by game and ones tested. This species is well suited to arid sites and livestock. provides an opportunity to enhance plantings where Plantings conducted within the Intermountain area few shrubs are adapted and most species are difficult exhibit adaptation to a wide array of sites. The species to establish. Desert ceanothus has been incorporated does express broad amplitude (Detling 1961). Of the into a few range and wildlife improvement projects in selections being tested, few exhibit adaptation to cal- central and southern Utah. careous soils common in Utah and Nevada. Plants Plant Culture—Seedlings establish best from late have established on these basic soils, but normally fall and winter seedings. Seeds are small, yet larger succumb in 4 to 6 years. Plantings established in than most other species of Ceanothus. There are ap- central Idaho on granitic neutral or slightly acidic soils proximately 23,000 cleaned seeds per lb (50,700 per have survived quite well. However, collections from kg) (Reed 1974). Seeds are borne in capsules that split central California and Oregon have suffered winter and disperse the seeds from the bush. Hot water damage. Normally portions of the crown are killed in treatments effectively improve germination. Untreated a single season; however, some plants have persisted seeds require approximately 42 days of wet prechilling for over 30 years. This species cannot be recommended to initiate germination (Stark 1966). for use on wildlands outside its natural range. How- Good seed crops are not produced each year, even ever, within its natural range it could be used for from bushes grown under cultivation. Generally, com- watershed protection and wildlife. It can be used for mercially harvestable crops are produced once every 3 low-maintainance landscaping in areas within its natu- to 4 years. Only about one-half the seeds acquired from ral range and on similar sites elsewhere. arid native stands and wildlife plantings in Utah are viable. Keeley (1977) reported similar numbers of

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 657 Chapter 23 Shrubs of Other Families viable seeds from wildland collections at other loca- separate from filled seeds using a gravity table or by tions. Plants flower in April and May; seeds ripen in flotation. July (Reed 1974; Van Dersal 1938) and can be hand Ecological Relationships and Distribution— collected by stripping the capsules. Like other species Deerbrush ceanothus is widely distributed in the Cas- of Ceanothus, desert ceanothus seeds disseminate as cade Mountains of Oregon and Washington south- they ripen. Seeds must be collected when slightly ward to southern California, Arizona, and New Mexico green and stored to allow drying. (Conard and others 1985; Kearney and Peebles 1960; Uses and Management—Desert ceanothus is a Munz 1974; Peck 1941). It grows as the dominant valuable browse species for big game (Bradley 1965; understory in a number of tree and shrub communities Gullion 1964; Leach 1956; Pase and Pond 1964; Powell (Conard 1987; Conard and others 1985; Gratkowski 1988), and is a particularly important winter forage 1961a; Hartesveldt and others 1975). It is also well (Conard 1987; Van Dersal 1938) and summer browse suited to open, dry slopes with and without shade for livestock (Boles 1987). (USDA Forest Service 1937). Based on its range of This shrub establishes quickly and is well adapted to adaptation and rapid growth, this species has been fresh disturbances. Horton (1949) reported that plants evaluated for watershed and wildlife habitat improve- survive well on road disturbances on coarse-textured ment projects in central Idaho (fig. 37). Plantings have soils at elevations near 5,350 ft (1,630 m) in California, established well from fall and winter seeding or and recommended planting container stock or direct transplantings of bareroot stock. Young plants grow seeding in spots to control erosion. Desert ceanothus rapidly; yearly growth rates have exceeded 30 inches recovers following fire because seeds stored in the soil (76 cm). In addition, the shrub exhibits an ability to are fire activated. Resprouting from root crowns does persist with understory grasses and herbs. The spe- occur, but only infrequently. cies sprouts aggressively following clipping. Although Varieties and Ecotypes—None. stem layering was reported by Biswell and Gilman (1961), this attribute has not been observed from plantings in Idaho. Deerbrush ceanothus grows well Family Rhamnaceae ______on fertile soils, but collections currently under study are not especially drought tolerant. Ceanothus integerrimus Deerbrush ceanothus Uses and Management—Collections from California and Oregon have been evaluated for their ability to Description—Deerbrush ceanothus is a spread- control erosion and restore wildlife habitat on log- ing, erect, deciduous shrub that grows rapidly ging roads and related disturbances where ground (Cronemiller 1959; Hitchcock and Cronquist 1973; cover is essential. The plants form extensive, fibrous Munz 1974; Van Dersal 1938). Leaves are large and root systems when nursery grown. The root system oblong-ovate in shape (Cronemiller 1959); some re- provides excellent soil stability, and plants effectively main on the bush throughout the winter (Cronemiller stabilize erodible slopes and unstable banks. Plantings 1959; Munz 1974; Van Dersal 1938). Like other spe- established at numerous sites in Idaho have been cies of Ceanothus, it is able to fix nitrogen (Biswell 1960; Harvey and others 1980). Plant Culture—Seeds ripen in midsummer and are dispersed as the capsule dries and dehisces (Keeley 1987). The seeds are smaller than those of most other Ceanothus species with about 70,000 cleaned seeds per lb (154,300 per kg) (Reed 1974). Seeds are long- lived and may persist in the soil for decades (Cronemiller 1959; Pase and Brown 1982; Quick and Quick 1961). Seedcoats are impermeable to water, and embryo dormancy also exists (Heit 1967a). Seed germination can be enhanced by heat pretreaments (Shmida and Barbour 1982), and is best induced by boiling the seeds for 1 to 3 minutes followed by wet prechilling (Quick and Quick 1961). Viability of deerbrush ceanothus seed lots usually is higher than for seed lots of other species of this genus. Over 85 Figure 37—Deerbrush ceanothus in south-central percent of cleaned seeds are often viable (Van Dersal Idaho plantings provide excellent wildlife habitat and 1938). Empty or immature seeds are difficult to watershed protection.

658 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families seriously weakened and thinned by cold winter tem- region has been limited. Plantings have been estab- peratures. The plants are not cold tolerant and serious lished on wildlife and watershed sites in the Interior dieback occurs each year. Less than 20 percent have Western States. Plant materials acquired from north- survived on these plantings. Winter losses have also ern California have been the primary source for most been noted from 1-year-old stock growing in nursery plantings. beds. The shrub has failed when planted on mine The species is best adapted to sites receiving 15 to disturbances and infertile sites in Idaho, Utah, Ne- 25 inches (380 to 640 mm) of annual precipitation. It vada, and Wyoming, although Biswell (1960) and has not proven adapted to basic soils, and it is not Harvey and others (1980) reported the plant is able to winter hardy. Less than 10 percent of all plants have fix nitrogen and has been successfully planted on survived for more than 5 years in plantings in the disturbed sites in the Southwest. It has not done well Intermountain region. throughout the Intermountain region on disturbances This species has received considerable use by big lacking topsoil. It does well on road fills created in the game. Its rapid rate of growth is an impressive feature. Idaho Batholith, but not on mine wastes. It is an attractive shrub, particularly when in bloom Although game animals utilize the shrub on re- (fig. 38). The flowers are abundant and cover the bush search plantings and the plant provides considerable with a dark blue blanket. Seed production is erratic. cover, it is not native to the Intermountain region, and Plants are easily cultivated and have considerable replacement of other shrubs with this species is not potential for horticultural uses. They can also be used advisable. It is not sufficiently winter tolerant to in low maintenance plantings and on recreational persist within the Intermountain region. sites. This species cannot be recommended for use outside its natural range. However, within its area of Varieties and Ecotypes—None. adaptation the shrub can and should be used to restore disturbed communities and provide wildlife forage Family Rhamnaceae ______and watershed protection. Ceanothus lemmonii Varieties and Ecotypes—None. Lemmon ceanothus Family Rhamnaceae ______Description—Lemmon ceanothus is a low, evergreen, spreading shrub that attains a height of 1.5 to Ceanothus prostratus 3 ft (46 to 91 cm); leaves are alternate, elliptical, 0.5 to Prostrate ceanothus 1.2 inches (13 to 30 mm) long, bright green and waxy above. Flowers are pale blue and abundant; seeds Description—Prostrate ceanothus is a low decum- form within capsules (Munz and Keck 1959; Sampson bent or prostrate, evergreen, layering shrub most and Jespersen 1963). prevalent in California (fig. 39). It grows 2 to 6 inches Ecological Relationships and Distribution— (5.1 to 15.2 cm) in height; stems root at the nodes, Lemmon ceanothus is a less widely distributed shrub than other species of Ceanothus. It is primarily confined to ponderosa pine communities from the base of the Sierra Nevada Mountains from Tuolumne and Eldorado Counties northward, and in the inner Coast Ranges from Lake and Yuba Counties north to Humboldt and Shasta Counties, California (Munz and Keck 1959). Although regionally adapted, it is an important deciduous shrub for wildlife (Sampson and Jespersen 1963). Plant Culture—Seeds of lemmon ceanothus have an impermeable seedcoat and embryo dormancy. Heat scarification reduces seedcoat impermeability and hastens germination. Following heat treatment, wet prechilling for approximately 2 months is required to relieve embryo dormancy. Young seedlings are vigorous and can survive some herbaceous competition.

Uses and Management—Research with this spe- Figure 38—Lemmon ceanothus is covered by cies for revegetation plantings in the Intermountain masses of tiny blue flowers in spring.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 659 Chapter 23 Shrubs of Other Families

field planted. If plants begin growth before field planting, they are susceptible to frost and drought. Seed- lings and young plants are vigorous but grow slower than most other ceanothus species. New seedlings and transplants survive well on disturbed soils (Brown and others 1971; Tiedemann and others 1976). Uses and Management—Prostrate ceanothus has been widely used for soil stabilization (Brown and others 1971; Tiedemann and others 1976; USDA For- est Service 1976), conservation (Plummer 1977), and horticultural plantings (Van Rensselaer and McMinn 1942), particularly in central California. Extensive transplanting projects have been employed to treat harsh, erodible sites. Prostrate ceanothus has attrac- Figure 39—Plants of prostrate ceanothus form a tive evergreen leaves and has been used to provide dense, low ground cover on an unstable granitic ground cover and serve as background plants to comple- slope in central Idaho. ment floral plantings and mixed shrub beds. Prostrate ceanothus has been successfully used to control erosion and stabilize roadways, and logging and mine disturbances in the Intermountain area. Its spreading to form dense mats 2 to 8 ft (0.6 to 2.4 m) ability to fix nitrogen (Delwiche and others 1965; across. Leaves are opposite, dark green, smooth above Stewart 1967) possibly contributes to its success on and sometimes gray green beneath. The margins usu- infertile soils. Collections from Oregon and California ally have coarse teeth near the tip. Flowers are blue have been widely used and are well adapted to moun- and borne in umbels. Fruits are roundish capsules tainous areas dominated by forested communities. (Sampson and Jespersen 1963). Most of these collections are tolerant of both shaded Ecological Relationships and Distribution— and open planting sites. The planting stock is widely Prostrate ceanothus occur in California on open flats adapted to infertile soils, exposed substrata, and mine and in pine forests at elevations from 3,000 to 6,500 ft wastes. Townsend (1966) reported this species recov- (910 to 2,000 m) from Calaveras and Alpine Counties ers quickly following fires, and is one of the first northward to Modoc County, and west to Siskiyou and species to reoccupy disturbances. Trinity Counties. It also occurs in Washington, Or- Prostrate ceanothus does not persist well with other egon, and western Nevada (Hitchcock and Cronquist plants. It spreads to provide a dense ground cover 1973). It grows in mountain shrub, ponderosa pine, intermixed with few other plants. Prostrate ceanothus and red fir forests of higher mountain regions, usually grows very slowly, and it spreads by stem layering. forming carpets (Sampson and Jespersen 1963). A Numerous shallow roots are formed along the spread- disjunct population occurs near Council, ID. ing stems. Prostrate ceanothus is able to persist and Plant Culture—Seeds are large, ranging from spread when buried by unstable soil. It is particularly 37,500 to 44,500 per lb (82,700 to 98,100 per kg). useful for stabilizing steep banks, roadcuts, and un- However, they are difficult to collect from the low- stable surfaces; however, it does grow slowly. Wildlife growing plants. Seeds must be hand collected by have made little use of the shrub on roadway plantings picking small groups of fruits before the capsules are in Idaho, which allows it to be used on steep sites fully ripe. When dry, the capsules dehisce and scatter where animals may cause damage. However, Sampson the seeds. and Jespersen (1963) reported that the shrub is browsed Seeds persist for many years in the soil, and can be heavily by deer in California. Kufeld and others (1973) stored under warehouse conditions for long periods reported, from a summarization of studies in Califor- without loss of viability. Seed germination is fire nia, that the species is moderately used in all seasons. dependent. Treatment of the seeds with boiling water This species provides attractive cover and has been for 0.5 to 3 minutes hastens germination (Reed 1974). used to landscape recreational sites and wildland Seedlings develop well from fall and late winter areas where low maintenance is provided. seeding. Varieties and Ecotypes—Various ecotypes and Nursery-grown bareroot stock establishes well as do collections of prostrate ceanothus have been devel- rooted cuttings. Container stock can be propagated oped for horticultural uses. Stock can be purchased from seed or from stem cuttings (USDA Forest Service from wildland nurseries. A collection acquired near 1976). However, transplants must be dormant when Council, ID, is particularly adapted to more arid sites,

660 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families surviving in areas receiving 15 to 20 inches (38 to 51 and bristlecone pine communities in Utah. Youngblood cm) of annual precipitation. It does well in open areas, and Mauk (1985) reported that Martin ceanothus and is better adapted to sites supporting other shrubs occurs in limited areas within the conifer forest types and some herbs than most other collections. of central and southern Utah. The plant is not highly shade tolerant, but is able to persist beneath Gambel oak, mountain maple thickets, and ponderosa pine Family Rhamnaceae ______overstories, and it responds quickly to burning or Ceanothus martinii other methods of clearing. It is noticeably absent from Martin ceanothus the aspen communities in Utah described by Mueggler and Campbell (1986), possibly due to the dense over- Description—Martin ceanothus is a low, rounded story provided by this tree species. Where it grows to spreading, unarmed shrub, between 8 and 32 inches with other shrubs, particularly Gambel oak, it is most (20 to 81cm) tall (Welsh and others 1987). The leaves prevalent in small, open areas where overstory species are alternate and deciduous with short petioles. Blades do not occur. are green on both sides, 0.3 to 1.2 inches (7.6 to 30.5 At lower elevations Martin ceanothus normally grows mm) long, 0.2 to 0.8 inch (5.1 to 20.3 mm) wide, elliptic as scattered plants with big sagebrush, skunkbush to oval, ovate, or obovate, and entire or serrulate sumac, antelope bitterbrush, and rubber rabbitbrush. except at the base. The inflorescence is corymbose Populations occur as somewhat isolated stands on with numerous white flowers (fig. 40). The fruit is a very harsh sites that also support green ephedra, three-lobed capsule 0.16 to 0.2 inch (4 to 5 mm) thick. Wyeth eriogonum, mountain ash, and bristlecone pine. Seeds are shiny brown and 0.1 inch (3 mm) long. It exists as a minor component in Gambel oak, creep- Sutton and Johnson (1974) reported the plant is ever- ing barberry, and gray horsebrush communities. It green, although plants may retain their leaves until seldom forms thickets. It often occupies ridgetops and late into winter when most are shed. exposed south and west slopes where few other species exist. It is well adapted to rocky, well-drained soils Ecological Relationships and Distribution— (Van Dersal 1938), and frequently occurs as the domi- Martin ceanothus is much more limited in distribution nant plant in these situations. than most other species of ceanothus in the Inter- This shrub is fire tolerant, but would not be consid- mountain region. It occurs in Utah, Nevada, Colorado, ered a seral species in most situations. Although it and Wyoming, at elevations between 6,000 and 9,500 recovers quickly following fire, it does not spread ft (1,800 and 2,900 m) (Welsh and others 1987). Kearney rapidly to occupy extensive areas. Other mountain and Peebles (1942) reported the plant grows in the brush species also recover well and are able to main- Kaibab Plateau and Grand Canyon areas at about tain their postfire position in these communities fol- 7,500 ft (2,300 m). lowing burning. Martin ceanothus grows in small patches or as scattered plants in pinyon-juniper, mountain brush, Plant Culture—Plants begin growth early in spring, big sagebrush, ponderosa pine, Douglas-fir, aspen, and leaves normally appear by early April. Flowers are formed in clusters on short stalks that often overtop the leaves and foliage. Under wildland conditions not all plants flower profusely each year. Good seed crops normally occur about once every 3 years, but even during periods of low production, some bushes produce considerable seed. Grazing by big game and livestock normally restricts seed production on many wildland sites. Plants established at protected nursery sites in Utah and Idaho have developed good seed crops nearly every year. Plants reach seed-bearing age in about 4 years if protected from grazing. Thereafter, annual seed production increases for about 3 years or until plants attain full stature. When plants are in full bloom, the bushes are very attractive and colorful. Flowering occurs in May and June, and seeds mature from mid-July to mid-August (Plummer and others 1968). Flowering generally occurs late enough in spring that seeds are not damaged. Figure 40—Martin ceanothus receives extensive use Flowers are insect pollinated and attract large by wildlife and livestock. numbers of insects. Developing seed crops are frequently

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 661 Chapter 23 Shrubs of Other Families damaged by insects, causing some to abort. Seeds are The shrub survives well on mine wastes and associ- formed in three-lobed capsules that are gummy and ated disturbances. It spreads slowly by natural re- sticky when green. As the fruit ripens, the capsules cruitment, but it is persistent and long lived. It is split open and seeds are dispersed 1 to 6 ft (0.3 to 1.8 m) adaptable to open slopes where big game may concen- from the plant. Rodents eagerly gather the ripening trate throughout the winter. The shrub has a low, fruits and cache the seeds. The seeds or fruits are rounded growth habitat; its forage is usually available difficult to harvest for commercial sale. Hand collec- to grazing animals. tion, a slow process, is required to remove the fruits Herbage quality is quite high, particularly for native from the bush. The fruits must be collected when stands in central Utah. Deer, elk, and livestock browse slightly green before the seeds are dispersed. When this species heavily (Diebert 1968). Tueller (1979), green, the fruits adhere tightly to the plant and are not however, reported only minimal use by big game in easily dislodged by beating or flailing. Once the fruit is Nevada. When planted on mine sites and on big game collected, it is air dried to allow the capsules to dehisce. habitat improvement projects in Utah, the plants are During drying, fruits should be covered with a light heavily browsed by big game and livestock. Use can screen to prevent the seeds from being thrown off the often be severe enough to reduce stand establishment. drying tables. Seeds can be separated from the debris Big game animals seek areas occupied by the shrub in using fanning mills or screens. After screening, empty late fall and winter. Both leaves and stems are grazed seeds can be removed on a gravity table or by floating during that season. the seeds in water. Cleaned seeds can be stored in open Seed production and harvesting are problems pre- warehouses for 10 to 15 years without loss of viability venting the wide use of this species. Plants grown (Plummer and others 1968). under protection and in nursery conditions produce Fresh collected seeds are dormant. Germination can abundant seed crops, but fruits are located amid the be aided by immersion in hot water. The most effective twigs in small clusters that are difficult to reach and treatment has been to place the seeds in boiling water detach. Plants are low growing and relatively uniform for 1 to 2 minutes. The water is then allowed to cool to in size and shape. Consequently, mechanical harvest- room temperature, and the seeds are removed and ers could be developed to dislodge and collect the dried. Seeds are then fall seeded or wet prechilled for fruits. If such equipment were developed, use of this approximately 30 to 90 days before germination begins. species would be greatly enhanced. Martin ceanothus seeds should be fall seeded in a Varieties and Ecotypes—Collections of Martin firm, weed-free seedbed. Seeds are relatively small, ceanothus from Sanpete County in central Utah have round, and easily planted. They should be planted proven well adapted to wildlife habitats throughout 0.25 to 1 inch (0.6 to 2.5 cm) deep. Rodents will gather the State. This ecotype is a good seed producer, and is planted seed; consequently, late fall seeding is recom- heavily browsed by big game. It appears well suited to mended. Seedlings appear early in the spring and are infertile sites, including mine disturbances. Small tolerant of frost. Young seedlings grow moderately seed production fields have been established, but seed well, but are not able to withstand competition from yields are not sufficient to meet current demands. herbaceous plants. Martin ceanothus should not be Commercial production should be encouraged. seeded directly with herbs, but in separate spots or strips. Seedlings are well adapted to harsh sites, and survive well from plantings on well-drained, infertile Family Rhamnaceae ______soils free of herbaceous competition. Transplants are easily cultured as bareroot nursery Ceanothus sanguineus stock or container material. Bareroot and container Redstem ceanothus stock must be dormant when field planted. If plants are lifted from the nursery bed and field planted after Description—Redstem ceanothus is an erect, de- vegetative growth begins, serious losses will occur. ciduous, nonrhizomatous shrub, 5 to 10 ft (1.5 to 3.0 m) tall (fig. 41); with purple or reddish, flexible, glabrous Uses and Management—Martin ceanothus is one twigs (Hitchcock and Cronquist 1973; Munz and Keck of the most promising and useful shrubs for rehabili- 1959). The leaves are thin, broadly elliptical to ovate tation of big game winter ranges. It is well adapted to or obovate, rounded or subcordate at the base, 0.4 to big sagebrush and mountain brush communities. It 3.9 inches (1.0 to 10 cm) long, subglabrous, and glan- occupies some of the most critical wintering areas in dular-serrulate (Munz and Keck 1959). The flowers Utah, including open south and west aspects, and it are numerous, white, and borne in showy compound has established well when planted in these situations. clusters 2 to 4 inches (5 to 10 cm) long from previous Seedlings are sensitive to drought and must be planted year’s wood (Kartesz and Kartesz 1980; Munz and in areas free of weeds. They grow at a moderate rate Keck 1959). The fruit is a slightly lobed, 3-celled capsule and can withstand heavy grazing even as young plants. about 0.2 inch (4 mm) long with one seed per cell.

662 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

following disturbances and becomes a dominant cover (Zamora 1975). Nearly closed stands may develop within 3 to 5 years (Morgan and Neuenschwander 1988). Like other species of Ceanothus, redstem ceanothus is primarily fire dependent for natural regeneration (Antos and Shearer 1980; Mitchell 1983; Wittinger and others 1977). Heat scarification stimulates seed germination (Lyon and Stickney 1976), and new seedlings flourish following wildfires or controlled burns. The species is capable of resprouting following burning and crown damage by deer (Miles and Meikle 1984). However, mature plants can disappear as shade increases. Periodic reburning maintains the species as Figure 41—Redstem ceanothus resprouts rap- an “obligate pioneer” (Laursen 1984). The shrub sur- idly following logging in central Idaho. vives with 10- to 15-year intervals between burning (Lyon and Stickney 1976). Plant Culture—Redstem ceanothus shrubs are normally robust and healthy plants, but abundant Ecological Relationships and Distribution— seed crops are infrequent. Individual plants may de- Redstem ceanothus is distributed from British Colum- velop heavy seed crops, and the better bushes are bia and Montana south to Washington, Oregon, Idaho, normally “high graded” during seed collection. Most and California (Hitchcock and Cronquist 1973; Munz plants are tall, and hand stripping is difficult. Fruits and Keck 1959; Reed 1974). This species is best adapted are not easily dislodged from the plant by beating or as an understory species in open or partially shaded shaking the stems. Seeds are dispersed as the capsule areas (USDA Forest Service 1937). It occurs at mid to ripens; thus, capsules must be collected when still low elevations to 2,400 ft (730 m) in Montana (Dittberner green and allowed to dry. Seeds are fully developed and Olsen 1983), and 4,000 ft (1,200 m) in California before the capsule dries; thus, seed quality or viability (Munz and Keck 1959). It is most abundant through- is not adversely affected by harvesting green capsules out much of the Pacific Northwest at mid elevations in shortly before they dry. Rodents actively gather the burned forest communities (Mueggler 1965). It is a ripening seeds (Conard and others 1985) and will principal component of extensive brush fields that collect entire seed crops as the fruits ripen. develop after fires, logging, or clearing (Halpern 1988). Seedbanks develop through annual accumulation The shrub is most prevalent in early seral stages of of seeds in the soil (Kramer 1984; Morgan and grand fir, Douglas-fir, western hemlock, and mixed Neuenschwander 1985). Although seeds are eagerly conifer forests (Armour and others 1984; Conard and eaten by rodents (Conard and others 1985), numerous others 1985; Dahlgreen 1984; Franklin and Dyrness seeds may accumulate in the soil and may remain 1973; Schoonmaker and McKee 1988). It also grows as dormant for decades (Furniss and others 1978). a scattered understory in mesic, open sites with pon- Seed densities ranging from 1.9 to 107 per ft2 (20 to derosa pine in central Idaho and the Northwest 1,150 per m2) have been reported in Idaho (Kramer 1984). (Franklin and Dyrness 1973). Miles and Meikle Large amounts of redstem ceanothus seeds have not (1984) reported that within the dry submontane been collected for commercial sales. The shrub usually Douglas-fir subzone, redstem ceanothus favors sites recovers satisfactorily by natural seedings so artificial with low evapotranspiration. planting is generally not necessary. However, seeds Redstem ceanothus is less shade tolerant than other are easily cleaned for storage or seeding. Seeds are species that coexist in brush fields of the Northwest approximately 0.08 inch (2 mm) in diameter; smaller (Hooker and Tisdale 1974). It is often replaced by than those of most Ceanothus species. There are about oceanspray and chokecherry as tree canopies close 130,000 cleaned seeds per lb (287,000 per kg). Seed (Conard and others 1985), yet its presence with other cleaning is quite simple. The capsules are allowed to shrubs provides extensive brush fields for 25 to 50 air dry, causing them to break apart and propel the years (Morgan and Neuenschwander 1988). Miles and seed. Consequently, the capsules should be covered Meikle (1984) reported successional dieoff as a result with a screen to prevent loss of seeds as fruits ripen. of combined factors including drought stress, severe Seeds and debris can be separated using an air screen frost, utilization, and shade. Although the density of fanning mill. this species may decline rapidly as shade increases Seeds have a hard, impermeable seedcoat and a (Antos and Shearer 1980), it reestablishes quickly dormant embryo (Heit 1967a; Kramer 1984). They

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 663 Chapter 23 Shrubs of Other Families require a heat treatment to open the seedcoat or hilar useful in seeding fresh disturbances where few other fissure (Furniss and others 1978). Immersing the seed species are initially able to establish. Under many in boiling water at temperatures between 190 and situations where little or no vegetation exists, redstem 212 ∞F (88 and 100 ∞C) (Heit 1967a; Niering 1981) for ceanothus can be seeded with herbs to provide a mixed 1 to 3 minutes, followed by cooling in the standing composition of plants. The shrub competes with other water, reduces seedcoat imposed dormancy. Heat- herbs without serious loss of young seedlings. treated seeds require wet prechilling for approxi- Transplants are easily grown as bareroot or con- mately 60 days to enhance germination (Heit 1967a; tainer stock. Bareroot plantings survive as well as Reed 1974). Radwan and Crouch (1977) reported ger- container seedlings and are much cheaper and easier to mination after 1 month of wet prechilling, but 4 months transplant. Transplanting is a useful method of estab- was the optimum period. Miles and Meikle (1984) lishing the shrub on selected sites and locations. Trans- suggested seeds collected from different aspects and plants grow rapidly and furnish a rapid-developing elevations have different optimum wet prechilling cover, particularly if plants are inoculated with nitro- periods. Seeds can be stored for many years in an gen-fixing organisms. Even when planted on mine or open warehouse, but may be damaged by chalcids. roadway disturbances, redstem ceanothus is an Gratkowski (1973) and Radwan and Crouch (1977) aggressive early developer. recommended storage at 37 to 69 F (3 to 21 C) to ∞ ∞ Uses and Management—Redstem ceanothus is reduce insect infestations. an important forage and cover plant for big game and Heat-treated seeds should be planted in fall in a other wild animals. In the Okanogan Valley of Wash- firm, well-prepared seedbed at depths between 0.25 ington and British Columbia, redstem ceanothus is and 0.5 inch (6.4 and 12.7 mm). Broadcast seeding on one of two browse plants that provide 60 to 70 percent a loose, rough seedbed or surfaces with considerable of the winter forage consumed by deer (Miles and litter also is successful. When seeding on a bare min- Meikle 1984). Although it provides 11 percent of the eral surface, seeds should be incorporated into the soil available winter forage, it supplies 52 percent of the and not left on the surface. Lyon (1971), and Youngberg herbage consumed by big game. and Wollum (1976) found that seedling survival by In many burned or cleared sites, redstem ceanothus natural recruitment was much higher following fall is often heavily grazed, and all annual growth is burning than spring burning. Most seedlings devel- utilized. This species is particularly important for use oping from spring burning emerged a year following by elk during the winter months (Kufeld 1973; Leege the burn; thus, they were subjected to considerably 1972; Leege and Hickey 1975). It is grazed in summer more herbaceous competition. Leege and Hickey (1975) by both deer and elk, although the amount and period found that the amount of precipitation received from of use may vary (Conard and others 1985; Keay 1977; May through August had a significant effect on seed- Keay and Peek 1980; Thilenius 1960). Game animals ling survival. are frequently attracted to burned sites where this Natural recovery following wildfire is often dramatic, species is abundant. Important wintering ranges are as seedlings are able to establish with little site or surface dependent on burning and logging to maintain this preparation. A considerable number of seedlings natu- shrub (Leege and Hickey 1975). rally succumb the first or second year after disturbance Redstem ceanothus is particularly palatable to wild- (Lyon 1971; Youngberg and Wollum 1976). However, life. It furnishes considerable herbage in areas where seedlings of this species are quite vigorous and can the plant is not abundant or the dominant species. persist on exposed sites. High seedling losses from Grazing animals seek and utilize the new growth natural seeding is to be expected as seedlings compete (Gaffney 1941; Leege 1968; McCulloch 1955; Thilenius for limited resources. Although less than 4 percent 1960). Scattered plants growing in riparian areas and may remain after two growing seasons (Orme and forest openings are selectively used at all seasons. Leege 1976), sufficient plants often survive to fully The age of the plant and seasonal period of grazing occupy the site. affect its palatability. Leege (1968) reported that new Natural thinning is not as evident where controlled growth is much more palatable than older twigs, and seedings are conducted. If sites are not overseeded, following burning, the nutritional value decreases little dieoff or thinning will occur. Once seedlings with age (Asherin 1973; Leege 1969; Leege and Hickey attain 1 to 2 years of age, they are often of sufficient 1971). However, Miles and Miekle (1984) did not size and vigor to persist with herbaceous competition. recommend burning to regulate or enhance the forage Redstem ceanothus is capable of nitrogen fixation value of this shrub. (Torrey 1978), which enriches the biomass and density The protein content of this plant during winter is of associated species. Thus, this shrub serves as an about 10.1 percent; this provides the protein level important nurse plant in many situations (Rose and necessary for deer maintenance (Miles and Miekle Youngberg 1981; Scott 1970). It has been particularly

664 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

1984). In addition, the average fiber content is a The flowers are white, heavily scented, very abun- relatively low 34.7 percent. Livestock also are at- dant, and borne in more-or-less corymbose clusters on tracted to plants with high nutritive quality and make short branches (Mozingo 1987; Sutton and Johnson considerable use of this species. 1974). The fruit is a three-lobed capsule, with a single Redstem ceanothus provides important cover for seed per locule. The seeds are small and dark brown or wildlife including big game, small mammals, and tan with shiny coats (Welsh and others 1987). birds. It is particularly important in winter. It is also The plant forms a deep spreading root system, and an important ground cover, and provides soil stability nitrogen-fixing actinomycetes occur on the roots on burned slopes where erosion potential is high. It (Hickey and Leege 1970; Stanton 1974). Snowbrush provides protection for streambanks and moist areas. ceanothus does not spread by layering or root suckering, Following fires it recovers quickly and provides ground but by resprouting from the crown. The plants have a cover and soil stability on areas where slumping and distinct cinnamon odor if the leaves or stems are surface runoff are common. crushed (Welsh and others 1987). Seedling establishment is very good when redstem Ecological Relationships and Distribution— ceanothus is planted on disturbances; consequently, Snowbrush ceanothus grows from the coastal ranges this species is utilized to seed abandoned roadway and of California north to British Columbia and eastward logging disturbances. Both direct seedings and to Alberta, Montana, South Dakota, and Colorado transplantings have been successful in conservation (Mozingo 1987; Reed 1974). Hitchchock and Cronquist plantings to control erosion. The species, however, is (1973) reported that C. v. var. hookeri occurs on the not well adapted to mine wastes, and has been slow to west side of the Cascade Mountains from northern invade road and logging disturbances where vehicle California to British Columbia. East of the Cascades, activity has occurred and soils are compacted. It has var. velutinus occurs in California, Nevada, Utah, been very useful for restoring riparian disturbances, Colorado, and South Dakota. In Utah, Wyoming, and and withstands livestock use. However, grazing of Colorado, the species is less abundant and restricted young plants can weaken developing stands. to more specific communities. Goodrich and Neese This shrub is able to establish with seeded herbs. It (1986) reported the plant occurs primarily at eleva- can be seeded with herbs or transplanted into new tions between 7,000 and 9,000 ft (2,100 and 2,700 m) grass/forb seedings. It can be used as a pioneer species in the Uinta and Blue Mountains and the west por- to restore harsh sites, and provides favorable areas for tion of the Tavaputs Plateau, where it is mostly other plants to establish. confined to mountain brush, ponderosa pine, and Varieties and Ecotypes—None. aspen communities. This shrub has a broad ecological amplitude and is considered a late seral species in some communities, Family Rhamnaceae ______and an early seral species in others. It is most often Ceanothus velutinus codominant with ponderosa pine and Douglas-fir. It Snowbrush ceanothus Description—Snowbrush ceanothus is a low- spreading, many-stemmed, evergreen shrub (fig. 42) (Countryman 1982; Welsh and others 1987; Zamora 1982) with olive-green to reddish brown bark. It attains heights of 3 to 9 ft (0.9 to 2.7 m) when growing in thickets or closed stands (Sutton and Johnson 1974). On open, windswept sites it may grow as low, rounded plants usually about 3 ft (0.9 m) tall; however, on fertile sites individual shrubs may be 12 ft (3.7 m) tall (Stanton 1974). Leaves are 1.2 to 2.0 inches (3 to 5 cm) long with three main veins and a glossy dark-green surface (Miles and Miekle 1984; Stark 1966). They are alternate, thick, pubescent to glabrous, ovate to elliptic, and very fragrant (Sutton and Johnson 1974; Welsh and others 1987). The leaves often curl in drought conditions; this conserves water by reducing evapo- Figure 42—Snowbrush ceanothus is a common un- transpiration (Miles and Miekle 1984). derstory shrub in many ponderosa pine communities.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 665 Chapter 23 Shrubs of Other Families is also commonly encountered in various mountain damage the shrub, particularly during years of low brush, antelope bitterbrush, and forested shrub com- snow cover when the shrubs are exposed. Although munities. It is a principal species in forested commu- plants may be killed to ground level, regrowth from the nities in Oregon and Washington (Franklin and crowns can usually occur. Fires can damage and kill Dyrness 1973) and Idaho (Steele and Geier-Hayes mature plants, but regrowth does occur, and stand 1987, 1992). As an early seral species, it occupies regeneration is often accomplished with controlled Douglas-fir, white fir, grand fir, ponderosa pine, and burns. lodgepole pine forests (Dyrness 1973; Franklin and Snowbrush ceanothus exists on a wide number of Dyrness 1973; Halpern 1989; Schoonmaker and McKee soil types and parent materials. It is able to grow on 1988). well-drained, coarse-textured soils (Stephens 1973), Although snowbrush ceanothus commonly grows on but water-holding capacity and nutrient availability dry, open hillsides (Weber 1987), and is considered to greatly affect the presence of this shrub (Miles and be shade intolerant (Stanton 1974), it will persist as an Meikle 1984). Soils that contain sufficient moisture to understory in forested communities for considerable sustain growth into July and August are important to periods. Increasing closure of the canopy and competi- its survival. Snowbrush ceanothus occurs on slightly tion affect removal of the shrub. The rate of succession acidic and neutral soils, and is particularly adapted to is regulated by soil moisture, and snowbrush ceanothus soils derived from granitic parent materials (Sutton can maintain a presence on xeric sites as an edaphic and Johnson 1974; Watson and others 1980; climax community (Miles and Miekle 1984). The shrub Zavitkovski and Newton 1968). may persist as a midseral species with Douglas-fir, but Plant Culture—Plant growth and flowering are if tree stands are somewhat open, it may remain as a regulated by elevation. Vegetative growth normally permanent understory (Conard and others 1985). begins by mid-April at lower elevations and as late as The species responds well to disturbances, particu- early June at higher sites (Noste and Bushey 1987). larly fires (Halpern 1988). High-intensity fires are often Flower development begins early in the season, often followed by the rapid recovery and postfire dominance by mid-April. At higher elevations flowering may of this plant, usually within 1 year, if an adequate occur in June or July. Seed maturation occurs from seedbank exists and climatic conditions favor estab- July to August (Plummer and others 1968; Reed 1974; lishment (Halpern 1989; Halvorson 1982). Seed germi- Stark 1966; Swingle 1939). nation is activated by burning (Halpern 1988; Marshall Plants usually produce a number of flowers each and Waring 1984; Mozingo 1987). This species is able to year, but viable seeds may not always mature. Seed compete with pioneering herbs. Snowbrush ceanothus production varies annually among wildland sites. Few is, therefore, a major component of brush fields and areas produce abundant crops each year, but some shrubby communities that develop after wildfires shrubs will generally produce seed. Young plants that (Gratkowski 1961b, 1978; Morris 1958). It is often a occur on open burn sites are not always better seed primary understory component on ponderosa pine sites. producers than those existing in mixed shrub/conifer It may exist as a long-lived seral species or a climax forests. Normally, some sites will bear commercially dominant. Along with antelope bitterbrush, it forms harvestable crops each year, but sites must be located important associations as a dominant climax shrub and inspected regularly. (Franklin and Dyrness 1973). Seed is harvested by hand stripping the fruits before Snowbrush ceanothus grows across a wide eleva- they mature or the capsules dry. As capsules dry, tional range from 3,500 to over 10,000 ft (1,100 to seeds shatter immediately. Capsules ripen irregularly 3,000 m) (Mozingo 1987; USDA 1937). Aspect affects on the bush and fruits must be collected when still the distribution of the species due to increasing evapo- green. However, seeds are fully ripe before the cap- transpiration demands on south and west slopes, sules are dry, and harvesting of slightly green fruits particularly in pioneering systems (Miles and Miekle does not diminish seed quality. Seed harvesting is 1984). The shrub is unable to establish on south, slow and expensive, and is a principal deterrent to the southwest, or west aspects at elevations below 3,250 ft wide use of this species. (990 m) in British Columbia because of xeric condi- Yields of 50 to 75 lb per acre (56 to 84 kg per ha) of tions. At high elevations it is not normally found on cleaned seed have been obtained from sites in Idaho. steeper east, northeast, or north aspects because of Rodents actively gather the fruits and seeds, and will poor shade tolerance and competition (Miles and Miekle remove much of the seed as it matures. Nearly com- 1984). plete removal of the seed by rodents has been reported Although snowbrush ceanothus is a fire-dependent by Conard and others (1985). pioneer species, successional dieoff is caused prima- Mature seeds have a hard impermeable seedcoat rily by drought stress due to competition. Shading, and a dormant embryo. Heat and wet prechilling are frost damage, and utilization may hasten the process necessary for germination (Mozingo 1987). Soaking (Miles and Miekle 1984). Severe frosts frequently

666 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families the seed in boiling water for 1 to 3 minutes normally ranges from 21 to 62 lb per acre (23.5 to 69.5 kg per ha) enhances germination. Seeds are then germinated in (Rose and Youngberg 1981; Tiedemann 1981; a cool, moist environment at 34 to 41 ∞F (1 to 5 ∞C) for Zavitkovski and Newton 1968). Nodulated plants can 63 to 84 days (Reed 1974). Germination is usually improve growth of associated species. Scott (1970) completed in about 14 days (Quick 1935). Seed germi- reported Douglas-fir trees growing in the open were nation is usually very high, exceeding 70 to 80 percent half the size of seedlings nourished by snowbrush (Reed 1974). Mozingo (1987) suggested that the opti- ceanothus. mum period of exposure to heat varies geographically Snowbrush ceanothus has been particularly valu- among ecotypes, but differences have not been noted able as a pioneer species for improving poor soils and in separate collections obtained from central Idaho. disturbed sites. It has been extensively used to reveg- Seeds can be stored for long periods without loss of etate logging roads, mine wastes, and other distur- viability. Good stands have developed from planting in bances. It is particularly adaptable as a pioneer spe- the fall at a depth of 0.5 to 0.75 inch (1.3 to 1.9 cm). cies capable of stabilizing and improving soil Seedings also establish well by broadcasting seeds on conditions that can then be used to enhance the a rough seedbed in the fall or early winter. Plantings establishment of associated species (Conard and oth- on bare mineral soil or amid surface litter are both ers 1985; Geier-Hayes 1987; Steele and Geier-Hayes successful if the soil surface remains wet throughout 1992). The shrubs improve soil fertility, which pro- the germination period. Miles and Miekle (1984) re- motes early establishment and growth of other plants ported that natural seedings that germinate in the (Franklin and Dyrness 1973; Steele and Geier-Hayes autumn, following late summer or fall burning, sur- 1987, 1992). vived much better than those that germinate in spring This shrub has been one of the most successful after spring burning. species used to revegetate roadways and unstable Snowbrush ceanothus seedlings establish well on slopes in the Idaho Batholith, where erosion control is fresh, open disturbances and compete favorably with needed during the year of disturbance. Plants estab- pioneering herbs. The species establishes very well lish quickly, and small seedlings or transplants grow from artificial seedings and is particularly adapted to rapidly to furnish excellent ground cover. The plant unstable sites. Sutton and Johnson (1974) and Miles can be seeded in mixtures with fast-developing herbs and Meikle (1984) reported young plants grow slowly, including western yarrow, fireweed, and Canada gold- but Scott (1970) found that nodulated seedlings pro- enrod, or it can be transplanted at various spacings to duced 2.5 times more dry weight than non-nodulated reduce wind erosion and runoff. plants. Results from plantings conducted in Idaho and Ecotypes with different growth habits have been Utah do not support reports that seedling growth is selected and used for erosion control. Plants obtained slow. In fact, plantings on forested roads and study from windswept sites have dense, low-growing growth sites are vigorous and aggressive. Seedlings and young forms, and have demonstrated adaptability to a wide plants develop well and are able to establish and range of sites. These collections maintain a low, dense, persist with herbaceous competition. However, con- leafy growth habit that provides excellent ground siderable differences between nodulated and non-nodu- cover. One-year-old bareroot stock normally develops lated plants were also noted on these sites. a crown spread of 18 to 30 inches (46 to 76 cm) during A large number of natural seedlings can be expected the year of establishment. to emerge on burn sites. Natural thinning normally Plantings established along roadways have per- occurs soon after emergence (Lyon 1971), but suffi- sisted following wildfires, and their extensive root cient numbers usually survive to provide an adequate system stabilizes roadfills. Regrowth occurs quickly stand. Winter frost, grazing, and lack of soil water and new shoots protect the soil surface. The evergreen contribute to seedling losses. growth habit of this shrub also aids in furnishing Snowbrush ceanothus is easily grown as bareroot or yearlong soil cover. The plant has been widely planted container stock, and transplanting is an excellent for erosion control in the West (Dietz and others 1980; method for stabilizing disturbances. Snowbrush Fessenden 1979; Monsen and Christensen 1975; ceanothus forms a symbiotic relationship with Strep- Plummer 1977; Stark 1966). tomyces ceanothii to fix nitrogen (Hickey and Leege Snowbrush ceanothus provides forage and cover 1970). These organisms form clusters of modified for game and wildlife. It is regarded as a primary short roots, not typical nodules (Furman 1959; Torrey winter forage for big game, but use varies considerably 1978). Seedlings grown for field plantings should be among geographic regions (Martinka 1976; Stanton inoculated with these nitrogen-fixing organisms. 1974; USDA Forest Service 1976). Undoubtedly, differences occur among ecotypes, and collections from Uses and Management—Snowbrush ceanothus is some regions are much more heavily grazed than a particularly important species for restoring and others. Miles and Miekle (1984) reported that enhancing soil fertility. Annual fixation of nitrogen

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 667 Chapter 23 Shrubs of Other Families snowbrush and redstem ceanothus are the most im- Varieties and Ecotypes—None. However, ecotypes portant winter browse species for mule deer in the with low growth forms, and selections that receive Okanogan Valley in British Columbia, and that 60 to heavy browsing have been identified. These selections 70 percent of the forage consumed in winter is supplied could quickly be advanced with more extensive seed- by these two shrubs. Plants in Idaho (Leach 1956), ing projects. portions of Montana (Klebenow 1965; Youds and Herbert 1988), and some collections from central Cali- fornia received moderate to heavy use. Within the Family Salicaceae ______Great Basin this species receives much less use by big Salix species game (Tueller 1979). Although plants are browsed in Willow winter, use is not excessive. Collections of this species have been assembled and field planted at various Introduction—There are about 300 species of wil- locations in Idaho. Collections from California, north- lows worldwide; about 70 species are native to North ern Idaho, and Oregon have received more use than America (Brinkman 1974i). Willows are constituents most Intermountain region accessions. Use also varies of many riparian zones in the Western United States. by season (Martinka 1976; McCulloch 1955) and among Their root systems stabilize streambanks, while stems big game animals (Kelbenow 1965; Peek 1974; Watson and leaves dissipate flood energy and catch sediments. and others 1980). Willows contribute to the formation of overhanging Snowbrush ceanothus withstands considerable banks that provide shade, cover, food, and travel winter browsing and is able to recover without corridors for a large number of vertebrates and inver- apparent loss in vigor. Snow cover and winter frost tebrates and microsites for establishment of many damage reduce both forage availability and animal plant species (Clary and McArthur 1992; Kovalchik use. Many sites are often covered with deep snow, and Elmore 1992; Thomas and others 1979b). and plants are unavailable for long periods (Miles Damage to many riparian ecosystems has resulted and Miekle 1984). from improper grazing practices, logging, road con- Cattle generally make little use of this shrub (Curtis struction, recreation, water impoundments, and other 1952). However, Miles and Miekle (1984) noted that human activities. Land management efforts have they selectively used it in late fall and winter, thus recently centered on restoration and maintenance of competing with big game for winter browse. Sheep and these dynamic ecosystems. Research and riparian goats browse new growth after fire (Stanton 1974). classification studies are rapidly providing knowledge Nutrient value varies among sites. In British Co- of willow ecology and guidelines for management lumbia, Miles and Miekle (1984) found snowbrush and (Hansen and others 1988a,b, 1995; Kovalchik 1987; redstem ceanothus were the only two native shrub Manning and Padgett 1995; Padgett and others 1989; species that provided maintenance level protein for Szaro and Patten 1986; Youngblood and others 1985). deer in the winter months. Digestible energy was Natural regeneration processes and techniques for higher for snowbrush ceanothus than for all other restoration of riparian communities have also re- native shrubs. The winter fiber content of snowbrush ceived recent attention. Improvement of degraded ceanothus was 26 percent, lower than any other shrub riparian areas requires that factors causing degrada- tested (Miles and Miekle 1984). This is due to the tion be addressed (Briggs 1995; Briggs and others retention of leaves on the bush. This provided a much 1994). Restoration efforts are unlikely to be successful higher amount of forage with higher nutritive content unless the causes of degradation are first mitigated. than that of deciduous shrubs (Miles and Miekle Most willow species are easily propagated; but little 1984). In contrast, Dittberner and Olsen (1983) de- information is available for many individual species. scribed protein and energy values for this shrub as General techniques for vegetative propagation and poor. Blank (1984) and DeByle and others (1989) production from seed are provided below. Specific suggested that fire increases nutritive content, but requirements, where known, are described in the Miles and Miekle (1984) concluded that the changes do plant culture chapter for each species. not support the use of fire for nutrient enhancement. The plant has considerable value in landscape and Vegetative Propagation—Most willows can be conservation plantings. Its growth habit, color, and propagated from hardwood cuttings. Techniques for evergreen features provide attractive cover. The plant harvesting and preparing cuttings are provided in is suited for extensive cultivation and will survive in Chapter 29 and in many books on plant propagation. organized plantings. In addition, it can be used in Stems of most willows contain dormant, preformed low-maintenance situations, and is particularly use- root primordia. These are formed during the first year ful for restoring disturbances where native species and quickly develop into roots when conditions are are desired. favorable (Haissig 1970, 1974). Upland species such as

668 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Scouler willow do not produce root primordia. They Willows may also be used to stabilize eroding root more slowly, and root initials appear only at the streambanks and other disturbances. Such practices base of the cutting (Densmore and Zasada 1978). as wattling, brush matting, and branch packing are Cuttings should be collected on or near the planting used to provide physical stability as vegetative cover site whenever possible. Released varieties of some develops (Gray and Leiser 1982; USDA Soil Conserva- willow species are available from commercial nurser- tion Service 1992). Live and dead plant materials in- ies, and can be used if restoration using local material cluding stems and branches are buried, planted, or is not an objective or if local material is unavailable placed on the soil to reduce soil movement, protect (Carlson 1992; Darris and Lambert 1993; USDA Soil erosion control structures, and enhance vegetation Conservation Service 1993a,b). development. Live willow cuttings used in these proce- Unrooted cuttings may be used on sites with favor- dures often root, providing dense root systems and cover. able planting conditions and long growing seasons. Propagation From Seed—Willows are dioecious; Dormant cuttings may be taken from fall through male and female flowers are borne on separate plants. early spring. Densmore and Zasada (1978) found that Inflorescences are catkins or aments that, depending cuttings taken in early spring rooted more readily on the species, appear before, with, or after the leaves than cuttings harvested in late fall. Rooted cuttings (Brunsfeld and Johnson 1985). The flowers are polli- may be required for species that do not produce pre- nated by bees. Fruits are capsules; each contains formed root primordia and for plantings on sites with numerous tiny, hairy seeds. Seeds of most species short growing seasons, rapidly declining water tables, ripen and are rapidly dispersed in early to midsummer or other factors that might slow establishment. (Brinkman 1974i). Dispersal may be timed to coincide Rooted and unrooted cuttings should be planted in with low water levels. Seeds are carried over long areas appropriate for each species. Cuttings or rooted distances by wind and water. Hairs on the seeds may stock should be planted at depths permitting the aid in dispersal; they may also function in the germi- developing root system to remain in contact with the nation process (Martens and Young 1992). Summer- water table throughout the summer (Busch and others dispersed seeds may be viable for as little as 1 week 1992). Plantings may be conducted in fall when water (Densmore and Zasada 1983). They are nondormant levels are low and appropriate planting areas exposed. and germinate rapidly, often in 24 to 48 hours, if This aids planters in placing the cutting base or root deposited on wet, exposed substrates. Germination system at an adequate depth and permits early-spring occurs rapidly over a wide range of temperatures. A root growth, but it also exposes cuttings or plants to requirement for exposure to light may ensure that spring runoff and flooding. Some plantings must be germination occurs in areas free of vegetative compe- completed after spring runoff. Streams with artifi- tition. The ability to germinate over a wide range of cially manipulated flows present special problems, as temperatures may compensate for the short period of root growth may not be rapid enough to advance with viability (Densmore and Zaszada 1983). Seed coats are a rapidly declining water table. transparent, and embryos contain chlorophyll. Conse- Survival and growth of willow plantings may require quently, germination and seedling development are that vegetative competition be removed or reduced at rapidly initiated under favorable circumstances. the time of planting. Neiland and others (1981) found Wide dispersal enables willows to act as pioneer or that competition with tall, dense grass stands adversely early successional species following flooding, burns, or impacted survival. Svejcar and others (1992) and other disturbances that create suitable microsites for Conroy and Svecar (1991) obtained greater second- germination. Continued rapid growth enables seed- year survival of Geyer willow cuttings planted on bare lings to develop root systems that remain within reach ground than on Nebraska sedge/Baltic rush sites even of the declining water table in summer, compensate though depth to the water table in midsummer was for the late period of germination, and withstand less on the Nebraska sedge/Baltic rush sites. They flooding during the succeeding spring. interpreted these results to represent an interaction of Some arctic and high-elevation willows such as biotic and abiotic factors on willow establishment. grayleaf willow mature seed in summer, but delay Large planting stock may be used when depressed dispersal until fall (Densmore and Zasada 1983; Zasada water tables or browsing are problems (York 1985). and Viereck 1975). These seeds exhibit conditional Planting poles or dormant stubs permit the base of the dormancy; they are capable of germinating at high cutting to be placed in the water table; foliage is temperatures, but not at the lower ambient tempera- produced beyond the reach of browsing animals, and tures of the season. Embryos of late-dispersing species cover and soil protection are obtained quickly. How- generally contain low amounts of chlorophyll. Wet ever, pole plantings are costly, and natural regenera- overwinter chilling of these seeds releases dormancy, tion is generally unlikely to occur unless the water permitting seeds to germinate soon after snowmelt table is restored or browsing curtailed.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 669 Chapter 23 Shrubs of Other Families and maximizing the period for seedling development inches (3 to 6 cm) long and 0.4 to 1.2 inches (1 to 3 cm) during the short growing season. wide with entire to shallowly toothed margins. Upper Willow seeds may be hand harvested once the cap- surfaces are green, and lower surfaces are glaucous sules begin opening, or recently dispersed seed may be with prominently raised veins. Catkins emerge and collected from drifts (Brinkman 1974i). Use of a com- mature with the leaves. Staminate catkins are 0.4 to mercial shaker has been suggested for harvesting 0.8 inch (1 to 2 cm) long. Pistillate catkins are whitish yellow poplar seed; the feasibility of using this system yellow and 0.6 to 2.4 inches (1.5 to 6 cm) long. Floral for collecting willow seed has not been assessed (Cech bracts are narrow, pale yellow to light brown, and and Keys 1987). However, because ripening within sparsely to densely hairy. Capsules are very loosely stands, plants, and catkins is uneven, it is often advan- arranged and borne on stipes 0.08 to 0.16 inch (2 to 4 tageous to harvest entire catkins just as the capsules mm) long. They are ovoid conic, 0.2 to 0.3 inch (5 to 8 turn from green to yellowish or as the first capsules mm) long, short hairy, and long beaked. Stigmas are begin opening (Brinkman 1974i; Martens and Young two lobed (Brunsfeld and Johnson 1985; Hitchcock 1992). Catkins can then be spread to dry in an enclosed and others 1964; Welsh and others 1987). area until the capsules open. With either method, it is Plants begin flowering when 2 to 10 years old; imperative that maturation be monitored closely. optimum seed producing age is 10 to 30 years (Hansen Seeds may be planted without cleaning, other than and others 1988a,b; Rawson 1974). Bebb willow flow- removal of twigs, leaves, and catkin branches. Mar- ers from April to July or August (Gill and Healy 1974; tens and Young (1992) suggested that hairs not be ISTA 1966; Viereck and Little 1972). Fruits ripen and removed from the seeds; their function is not fully seeds are dispersed from May to August (Densmore understood. However, the hairs do cause the seeds to and Zasada 1983; ISTA 1966). cling together. They may be removed by gentle carding Ecological Relationships and Distribution— (Atchley 1989) or by forcing air through a series of Bebb willow is widespread in Canada and the United sieves containing the seeds (Fung and Hamel 1993; States, occurring from low to upper-middle elevations Harder 1970). Both techniques are rather slow if large (Brunsfeld and Johnson 1985; Hitchcock and others quantities of seed are to be cleaned. 1964). It does not occur west of the Cascade Mountains Brinkman (1974i) reported willow seed could be or in California. At low elevations Bebb willow grows stored for only 10 days at room temperature, but in broad meadows and on alluvial terraces and viability could be maintained for 1 month if seed is subirrigated slopes. It is most common on organic or stored wet in sealed, refrigerated containers. Zasada mineral soils with textures ranging from silty to sandy and Densmore (1977, 1980) found that seed of several or gravelly (Marchant and Sherlock 1984), and it will summer and fall-dispersing willows could be stored tolerate moderately alkaline to moderately acidic soils for up to 3 years if dry seed was placed in doubled 3-mil (pH 5.5 to 7.5) (Haeussler and Coates 1986). Water polyethylene bags and stored at 14 F (–10 C). ∞ ∞ tables are usually within 3.3 ft (1 m) of the soil surface Seeds may be sown directly into containers or onto throughout the summer. Mottled or gleyed soils are wet soil surfaces. In the bareroot nursery, seed may be common in the first 3.3 ft (1 m) (Hansen and others covered lightly using a roller (Brinkman 1974i). It is 1995). At upper elevations Bebb willow is found in essential that the soil surface be kept wet and shaded, drier riparian areas. if possible. Seeds may be broadcast on appropriate Uplands associated with Bebb willow may be domi- sites if surfaces remain wet. Little information is nated by Wyoming big sagebrush, mountain big sage- available on production of planting stock from seed or brush, Douglas-fir, or Engelmann spruce communi- direct seeding. Use of seed may be advantageous if ties. Aspen, black cottonwood, and water birch are local cutting material is not available. The ability to common overstory associates (Brunsfeld and Johnson store seed for up to 3 years provides some flexibility in 1985; Padgett and others 1989). planning propagation and in timing direct seeding. Bebb willow may dominate early seral willow communities along rivers and streams, overflow channels, Family Salicaceae ______or seeps (Boggs and others 1990). Seeds germinate on wet mineral substrates with full exposure to sunlight. Salix bebbiana Suitable seedbed microenvironments include recent Bebb willow, beaked willow alluvial deposits along streams and abandoned, silt- filled beaver ponds (Hansen and others 1988a). Seed- Description—Bebb willow is a thicket-forming lings colonize wet, exposed mineral soils following shrub or small tree growing to 33 ft (10 m) in height. fires and have been noted to invade mine spoils in Roots are shallow and dense. Twigs are reddish or southwestern Canada (Marchant and Sherlock 1984; grayish brown and hairy when young. Leaves are Watson and others 1980). Burned or otherwise dam- elliptic, obovate or oblanceolate, mostly 1.2 to 2.4 aged trees resprout readily. Plants may also develop

670 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families from detached branch segments (Watson and others the short-lived seeds to germinate in highly variable 1980). environments during the short Alaskan growing sea- Bebb willow communities are relatively stable once son. By contrast, freshly harvested Montana seeds established. They may persist for long periods due to were highly germinable at 59 to 77 ∞F (15 to 25 ∞C) and the presence of high water tables or short periods of less germinable at 41 to 50 ∞F (5 to 10 ∞C) (Atchley flooding (Hansen and others 1988a,b). Bebb willow 1989). Thus, germination may be inhibited by cool requires full sun and lacks shade tolerance (Watson seedbed microenvironments. Bebb willow seedlings and others 1980); it may be replaced by taller species. can tolerate some shade (Atchley 1989). Plant Culture—Bebb willow may be propagated as Uses and Management—Watson and others rooted cuttings, but reports of rooting capability vary. (1980) described Bebb willow as being a rapid grower, In Alaska, cuttings harvested in autumn after leaf fall capable of recovering rapidly after browsing, moder- or in spring following leaf expansion failed to root ately aggressive on suitable sites, a relatively good soil (Densmore and Zasada 1978). In contrast, rooting stabilizer, and exhibiting low drought tolerance. Stands success of current year’s softwood, also harvested in stabilize streambanks, providing protection from Alaska, was 42 percent (Halloway and Zasada 1979). flooding. Bebb willow often occurs in clumps rather In Montana, Atchley (1989) obtained 40 percent root- than continuous canopies (fig. 43). Thus, it provides ing when cuttings of 4-year-old wood were harvested shading for fisheries while permitting good access to in May. Platts and others (1987) reported dormant recreational fishing sites. Campgrounds cannot be cuttings rooted readily. Growth was initiated in about established within Bebb willow sites because the soils 10 days and shoot growth in 10 to 20 days. Roots were are generally wet early in the season. formed along the entire length of the stem. For production of rooted cuttings in Saskatchewan, Cram (1976) recommended that 6-inch (15-cm) cuttings be harvested in early fall and immediately placed in cold storage at 34 ∞F (1 ∞C) for spring planting in a bareroot nursery. Rooted cuttings were subsequently outplanted as 1-0 stock. Watson and others (1980) reported 30 to 70 percent survival of 1-0 rooted hardwood cuttings on mine overburden in northern Alberta; most mortality resulted from rodent damage. Unrooted cuttings are sometimes used. Hansen and others (1988a,b, 1995) recommended use of early spring- harvested dormant cuttings 12 to 20 inches (30 to 50 cm) long with diameters greater than 0.4 inch (1 cm). Bebb willow seeds must be harvested immediately when mature; seeds are dispersed in a period of 1 or 2 days (Marchant and Sherlock 1984). Fruits are green when preripe and yellowish when ripe (Watson and others 1980). Seeds may be hand harvested from trees or from recently dispersed drifts. Seeds may be used with or without cleaning (Brinkman 1974i). Atchley (1989) found that carding was more effective than the air-blowing technique described by Harder (1970) for removing the dense hairs from the seeds. Seeds vary in size and shape. Atchley (1989) found that viability was greater for plump, green seeds compared to small dark-brown seeds or wrinkled seeds. There are about 2,500,000 cleaned seeds per lb (5,500,000 per kg) (Brinkman 1974i). Bebb willow seeds collected in Alaska can be stored dry for at least 3 years in doubled 3-mil polyethylene bags at 14 ∞F (–10 ∞C) (Zasada and Densmore 1980). Fresh seeds collected in Alaska are nondormant. Germination at temperatures from 41 to 77 ∞F (5 to Figure 43—Thickets of Bebb willow dominate many 25 ∞C) exceeded 90 percent (Densmore and Zasada 1983; riparian communities from low to upper mid eleva- Zasada and Viereck 1975). This adaptation permits tions in western mountain ranges.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 671 Chapter 23 Shrubs of Other Families

Bebb willow provides a major source of highly pro- leaf surfaces are moderately pubescent when young. ductive and palatable browse for moose, deer, rabbits, Upper surfaces of mature leaves are green; lower elk, beaver, and bighorn sheep (Marchant and Sherlock surfaces are paler, but not glaucous. Catkins expand 1984; Watson and others 1980). It is more tolerant of with or slightly before the leaves. Staminate catkins browsing than Geyer willow, Booth willow, yellow are 0.4 to 1 inch (1 to 2.5 cm) long; pistillate catkins are willow, and Drummond willow, and may increase at 0.8 to 1.6 inches (2 to 4 cm) long. Floral bracts are their expense following heavy browsing (Hansen and brown to black and pubescent with long, curly, tangled others 1995). Use of Bebb willow increases in late hairs. Capsules are borne on stipes less than 0.08 inch winter. It remains accessible in deep snow as branches (2 mm) long. They are 0.12 to 0.24 inch (3 to 6 mm) are bent down and become accessible to browsing long, oblong ovate, and glabrous. Stigmas are nearly animals (Dorn 1970; Viereck and Little 1972). Song entire (Brunsfeld and Johnson 1985; Hitchcock and birds and game birds use the species for nesting and others 1964; Welsh and others 1987). food. Ecological Relationships and Distribution— Bebb willow communities are often adjacent to Booth willow occurs at midelevations from British meadows grazed heavily by livestock (Boggs and oth- Columbia to Alberta and south to northeastern Cali- ers 1990; Dorn 1970). Trees are usually widely spaced, fornia and Colorado (Brunsfeld and Johnson 1985). It allowing for easy access (Tesky 1992). Consequently, may dominate or codominate early seral willow com- deferring grazing until sites are dry may be required munities in fens, bogs, abandoned beaver ponds, wet to reduce livestock trampling and soil compaction meadows, and riparian sites, ranging from recently (Boggs and others 1990). Heavy livestock use may exposed rocky or gravelly deposits with high water restrict Bebb willow seedling establishment, resulting tables to drier benches with deep sandy to clay-loam in stand degeneration. soils and water tables to 3.3 ft (1 m) below the soil Prescribed burns can be used to rejuvenate de- surface (Brunsfeld and Johnson 1985; Manning and graded Bebb willow stands (Hansen and others 1988a,b; Padgett 1995; Padgett and others 1989; Youngblood Tester and Marshall 1962). Burned plants sprout and others 1985). Booth willow will tolerate moder- rapidly from basal stems. The small seeds are dis- ately alkaline, but not strongly acidic or basic soils persed over considerable distances (Haeussler and (Haeussler and others 1990). Coates 1986; Kovalchik and others 1988). Establish- Booth willow often grows in association with Geyer ment from seed depends on timing of the fire and willow and whiplash willow in open bottom lands availability of wet mineral soils to support germina- (fig. 44) and with Drummond willow in forested tion and seedling establishment (Chrosciewicz 1988; stream bottoms. Understory species range from Viereck and Little 1972; Viereck and Schandelmeirer beaked sedge and Nebraska sedge in wetter commu- 1980). nities to various forbs and shrubs in drier areas. Bebb willow is used as a “diamond willow” for carving Kentucky bluegrass may be a dominant understory canes, lampposts, and furniture (Watson and others 1980). Wood of this species has also been used for baseball bats, charcoal, and gunpowder (Viereck and Little 1972) . Varieties and Ecotypes—“Wilson” Bebb willow was released in 1985 by the Alaska Plant Materials Center (Wright 1989). It exhibits wide adaptability and a dense growth habit; it is useful for windbreaks, living fences, and screens.

Family Salicaceae ______Salix boothii Booth willow Description—Booth willow is a highly branched, rounded shrub with numerous basal stems 10 to 20 ft (3 to 6 m) tall. Young twigs are yellow to dark brown and hairy, becoming glabrous at maturity. Mature leaves are elliptical to broadly lanceolate, thick, firm, and about 0.8 to 3 inches (2 to 8 cm) long. Margins are Figure 44—Shrubby Booth willow is overshadowed by entire to toothed with fine, gland-tipped teeth. Both treelike whiplash willow.

672 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families species in areas with histories of grazing distur- dominant in Booth willow communities, is generally bances and dry surface soils (Hansen and others more palatable. Livestock use of both species should be 1988a,b). Uplands associated with Booth willow restricted until soils are dry to avoid compaction support sagebrush, juniper, western spruce-fir, (Hansen and others 1988a,b). Production of Booth lodgepole pine, Douglas-fir, and alpine meadow com- willow stands is often high, while understory produc- munities (Esser 1992). tion is variable, depending on the amount of willow Booth willow resprouts following burning or me- canopy present. Heavy use on sites with fine soils and chanical damage. Resprouting is more vigorous if stem palatable understory species must be avoided to pre- removal occurs during the dormant season (Kovalchik clude soil erosion and bank sloughing. and others 1988). Booth willow is also tolerant of frost Prescribed burning may be used to rejuvenate deca- and flooding. Efficiency of gas exchange and root dent stands and increase sprouting for use by big game regeneration are increased by the presence of aeren- (Haeussler and others 1990; Manning and Padgett chyma; adventitious rooting may occur above the level 1995). Plants also reestablish from offsite seed sources of flooded soil (Kovalchik 1992). However, growth is if burning occurs prior to seed dispersal and fires are reduced by prolonged flooding above the level of the severe enough to expose mineral seedbeds (Densmore root crown. and Zasada 1983; Zasada and Viereck 1975). Flowering occurs from May to June. Fruits ripen Varieties and Ecotypes—None. and seed is dispersed in late July and early August (Haeussler and others 1990; Zasada and Viereck 1975). Greatest quantities of seeds are produced by plants Family Salicaceae ______that are 2 to 10 years old (Brunsfeld and Johnson 1985; Haeussler and others 1990). Booth willow regenerates Salix drummondiana from short-lived seeds that are dispersed by wind or Drummond willow, beautiful willow, blue water. Suitable microsites for germination include willow wet sand or gravel surfaces receiving full sunlight (Hudak and Ketcheson 1992). Description—Drummond willow is a multiple- stemmed shrub usually 10 to 13 ft (3 to 4 m) tall. Young Plant Culture—Booth willow may be used to reveg- twigs are finely hairy and yellow or green, later becom- etate degraded riparian areas. Cuttings root readily ing glabrous and purple brown to yellow with a bluish and abundantly along the lower one-third of the stem glaucous bloom. Mature leaves are elliptic to lan- (Platts and others 1987). New roots and stems are ceolate or oblanceolate, 1.6 to 3.5 inches (4 to 9 cm) long initiated in 10 to 15 days. Seedlings must be protected and 0.5 to 0.8 inch (1.3 to 2 cm) wide. Margins are from browsing for at least 3 years to ensure establish- entire and somewhat revolute. Leaf surfaces are dark ment (Kay and Chadde 1992). green above and glaucous to strongly silvery pubes- Fresh seeds harvested in Alaska germinated rapidly cent beneath. Catkins expand before or with the leaves; and nearly completely at constant temperatures flowering generally occurs in May (Dittberner and ranging from 41 to 77 ∞F (5 to 25 ∞C) (Densmore and Olson 1983; Munz 1973). Staminate catkins are 0.4 to Zasada 1983). This behavior may be an adaptation to 1 inch (1 to 2.5 cm) long. Pistillate catkins are 0.6 to 1.6 a short period of viability, the short growing season, inches (1.5 to 4 cm) long, densely flowered, and silvery and extreme variability in microsite conditions. pubescent. Floral bracts are black or dark brown, long Uses and Management—Booth willow provides hairy, and persistent. Capsules are 0.12 to 0.16 inch browse for moose, elk, deer, small mammals, birds, (3 to 4 mm) long and densely short hairy. Stigmas are and other wildlife (Chadde and Kay 1988; Manning lobed (Brunsfeld and Johnson 1985; Hitchcock and and Padgett 1995; Youngblood and others 1985). Ex- others 1964; Sutton and Johnson 1974; Welsh and cessive browsing can reduce Booth willow regenera- others 1987). tion and seed production (Kay and Chadde 1992). Ecological Relationships and Distribution— Booth willow communities provide cover for many Drummond willow occurs from Yukon and terrestrial species and shade and overhanging banks Saskatchewan south to California and New Mexico for fish (Brunsfeld and Johnson 1985; Manning and and east across southern Canada and the Northern Padgett 1995). Streams in Booth willow stands may United States. It occurs at mid to high elevations from provide recreational fishing, but access is restricted by the upper sage to the spruce-fir zone (Brunsfeld and the dense canopies. Johnson 1985; Hansen and others 1995; Manning and Palatability of Booth willow is fair for livestock and Padgett 1995). It is found around springs, seeps, sheep; use increases late in the season (Manning and beaver ponds, lakes, ponds, and on flood plains and Padgett 1995; Padgett and others 1989; Youngblood and benches adjacent to swift, rocky, high gradient streams others 1985). Geyer willow, which is often abundant or (fig. 45). At the lower edge of its range it generally

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 673 Chapter 23 Shrubs of Other Families grows in wet, sandy or gravelly soils adjacent to Plant Culture—Drummond willow is useful for streams. In cooler habitats at higher elevations it is stabilization of disturbed sites adjacent to streams more abundant and occurs across broad valley bot- and ponds. The species may be established from toms. In such areas it grows on moist, well-aerated unrooted or rooted cuttings. Unrooted cuttings are mineral soils ranging from sandy to clayey loams. The most successful if planted on sites that remain wet water table may be near the surface in spring, but it throughout the growing season. Two to 4-year-old drops below 3.3 ft (1 m) by late summer. Common hardwood cuttings harvested in early spring root associates are Booth willow and Geyer willow. Due to readily with root initials appearing along the length of the similarity of their ecological requirements, com- the stem. Roots and shoots appear about 10 days after munity types in Utah, western Wyoming, and eastern planting (Platts and others 1987). Seed biology of Idaho dominated by either Drummond willow or Booth Drummond willow has not been studied. Seeds ripen willow are classified as Booth willow types (Padgett in summer and remain viable for about 1 week (Uchytil and others 1989; Youngblood and others 1985). How- 1991a). ever, Goodrich (1992) noted that the dominance of this Uses and Management—Drummond willow com- species and absence of Booth willow along rocky, high- munities are long lived, but shade intolerant. Produc- elevation streams of the Uinta Mountains indicates a tivity is generally high. Stands provide stream stabi- need to describe one or more riparian community lization, dense shade for fish, and habitat for beaver, types in which Drummond willow is the dominant birds, and other wildlife. woody species. Moose and elk make heavy use of Drummond willow, particularly in winter (Peek 1974; Singer 1979; Stevens 1970). Plants in Yellowstone National Park have been stunted by moose and elk use (Chadde and Kay 1988; Patten 1968). In Oregon, Drummond willow is considered highly palatable to livestock, big game, and beaver (Kovalchik and others 1988). Heavy browsing by big game may lead to degradation or loss of this species. Loss of willows resulting from heavy use by beaver may result in an increase in graminoids, loss of beaver populations, and lowering of the water table. Drummond willow communities stabilize streambanks and limit livestock access (Hansen and others 1988a,b). Heavy livestock use or trailing at streamside, particularly when soils are wet, however, may lead to highlining, dead clumps, stand loss, and bank compaction and sloughing. Decadent willows can recover rapidly if browsing is reduced. Excessive livestock grazing may also lead to a loss of palatable graminoids and their replacement by less palatable weedy species. Drummond willow stands may act as natural firebreaks in wet years, but during dry seasons they may burn (Crane 1982). Decadent Drummond willow stands may be rejuvenated using prescribed burning in late summer or early fall when sites have dried. Burning near streambanks should be carefully regulated. Livestock grazing must be curtailed for at least 2 or 3 years following burning. Plants recover by resprouting from the root crown following fire (Boggs and others 1990; Kovalchik and others 1988). Seed dispersed from unburned areas may germinate on exposed, wet mineral soils (Viereck 1982; Viereck and Schandelmeier 1980). Recovery from seed depends on seed availability, season of burning, and the presence of open, wet mineral soils. Varieties and Ecotypes—“Curlew” Drummond Figure 45—Drummond willow borders a high willow was cooperatively released by the USDA Soil gradient stream in eastern Utah. Conservation Service Pullman Plant Materials Center

674 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families in 1993 (USDA Soil Conservation Service 1993a). Patton 1987; Youngblood and others 1985). It also Curlew is valuable for inclusion in conservation grows on rocky, gravelly, and sandy lake and pond plantings for streambank stabilization, erosion con- edges; wet, well-drained alluvial terraces and bottom trol, improvement of wildlife habitat, naturalized land- lands that may have fine-textured soils (fig. 46); along scaping, and shelterbelts. It grows on wet, sandy to irrigation ditches; and in wet areas along roadways gravelly sites in areas receiving 20 to 25 inches (51 to (Brunsfeld and Johnson 1985). Soil surfaces in com- 64 cm) of precipitation. The release originates from a munities at the water’s edge are often characterized by native population growing near the Curlew River, high percentages of exposed areas and rocky surfaces, Ferry County, Washington. It is propagated from seed and provide seedbeds for establishment of a wide or cuttings. Hardwood cuttings taken in late winter array of annual and perennial herbaceous species are most commonly used. (Youngblood and others 1985). Sandbar willow may continue to dominate repeatedly disturbed areas, regenerating by rootsprouting or establishing from seed. Family Salicaceae ______Stabilized communities of this shade-sensitive species Salix exigua may eventually be replaced by cottonwoods or other Coyote willow, dusky willow, narrowleaf willow species. Two subspecies of coyote willow are widespread in willow, sandbar willow the Intermountain area; considerable variability oc- Description—Coyote willow is a colonial shrub curs within each subspecies. Salix exigua ssp. exigua that spreads underground to form dense thickets. occurs primarily in the Wyoming big sagebrush zone of Stems develop from shoot buds on lateral roots. Stems foothill areas and is distributed from southern British are numerous, slender, and up to 16 ft (5 m) tall. Young Columbia and Alberta to northern Mexico. This sub- twigs are thinly to densely hairy; older ones are gla- species is characterized by entire or few-toothed, brous and brown to reddish brown. Mature leaves are gray-green to silvery pubescent leaves; late-develop- linear to oblong, entire to minutely toothed, 0.8 to 4.3 ing catkins; and narrow, acute floral bracts. S. e. ssp. inches (2 to 11 cm) long, and 18 to 30 times longer than melanopsis is distributed from Alberta to California wide. Leaf surfaces are gray green to silvery white, and south to northern Colorado, and occurs at higher and glabrous to densely pubescent. Catkins terminate elevations than ssp. exigua (Hitchcock and others the twigs and emerge and mature with or after the 1964). Subspecies melanopsis is distinguished by its leaves. Staminate catkins are yellowish and 0.6 to 1.8 toothed to subentire leaves that are glaucous beneath, inches (1.5 to 4.5 cm) long. Pistillate catkins are 0.6 to glabrous capsules, and blunt floral bracts (Brunsfeld 2.4 inches (1.5 to 6 cm) long. Floral bracts are yellowish and Johnson 1985). and hairy. Capsules are 1.6 to 2.8 inches (4 to 7 cm) In addition to spreading by suckering, coyote willow long, sessile, and glabrous to hairy. Stigmas are deeply produces numerous small seeds that are dispersed by lobed (Brunsfeld and Johnson 1985; Hitchcock and wind and water (Arno and Hammerly 1977; Mozingo others 1964; Welsh and others 1987). Flowering occurs from April to July (Dittberner and Olsen 1983). Fruits mature and seeds are dispersed from May to July (Brinkman 1974i; Noble 1979; Ware and Penfound 1949). Seed dispersal may correspond to low water levels (Youngblood and others 1985). Ecological Relationships and Distribution— Coyote willow is widespread in North America, but occurs entirely east of the Cascade Mountains at low, or occasionally midelevations from the sagebrush to the spruce-fir zone. Coyote willow is one of the earliest pioneer species to colonize freshly deposited sand and gravel bars, which may be below the high water mark where it is subject to annual flooding, ice jams, and associated scouring and deposition (Hansen and others 1988a,b; Wasser and Hess 1982). It frequently occurs along streams and rivers in bottom lands, forming a narrow zone between the water’s edge and adjacent cottonwood, water birch, thinleaf alder, or Figure 46—Coyote willow forms a dense thicket other willow communities (Kovalchik 1987; Szaro and along a dry streambed.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 675 Chapter 23 Shrubs of Other Families

1987). Seeds germinate within 24 hours if exposed to Conservation Service, Pullman Plant Materials Cen- freshly deposited, wet alluvium in full sunlight ter in 1993 (USDA Soil Conservation Service 1993b). (Densmore and Zasada 1983). In the field, seeds gen- The release originated from material collected on the erally do not remain viable for more than 1 week (Ware Tucannon River, near Starbuck, Washington, at an and Penfound 1949). Branches are flexible and resprout elevation of 560 ft (170 m). It is recommended for use if buried in sediment. Plants may also regenerate in conservation plantings for riparian area stabiliza- vegetatively from broken stems and roots deposited on tion and restoration, wildlife habitat improvement, exposed, wet soils. and shelterbelts. It is normally propagated from cuttings, but seed can be used. Plant Culture—Coyote willow is an excellent species for restoration of disturbed riparian areas and recently deposited sediments because of its early suc- Family Salicaceae ______cessional status, rapid growth, and ability to root sprout (Uchytil 1989a). Unrooted cuttings may be Salix geyeriana used successfully on low-elevation sites where the Geyer willow water table remains high throughout the growing season. Rooted stock should be used on sites with Description—Geyer willow is a large shrub or fluctuating water tables, high flooding potentials, or small tree with numerous slender, ascending stems 10 short growing seasons. Coyote willow cuttings develop to 13 ft (3 to 4 m) in height arising from a tight basal roots along the entire length of the stem (Platts and cluster (fig. 47). The root system is deep, fibrous, and others 1987). Roots and shoots appear in about 10 spreading (Sutton and Johnson 1974). Young twigs days. are densely hairy, but become glaucous; older twigs There are about 10,000,000 cleaned seeds per lb (22,000,000 per kg) (Brinkman 1974i). At 72 ∞F (22 ∞C), Brinkman (1974i) obtained 83 percent germination in 4 days. Uses and Management—Maintenance of coyote willow stands is important because of their ability to stabilize streambanks (Uchytil 1989a). Loss of this species as a result of browsing and human-caused disturbances can lead to serious erosion problems (Hansen and others 1988a,b). Coyote willow is browsed by moose, elk, and to a limited extent, mule deer (Patten 1968; Van Dersal 1938). It is heavily used by beaver (Mozingo 1987). Thickets of sandbar willow provide valuable cover for birds and other wildlife species (Lindauer 1983). How- ever, because of their upright growth habit, they provide only moderate amounts of shade for fish (Hansen and others 1988a,b). Dense stands of coyote willow bordering streams may limit access by fisher- men. As such stands are seasonally flooded, they should not be used for campgrounds or roadways (Hansen and others 1988a,b). Coyote willow is used for forage and especially for cover by livestock (Van Dersal 1938). Forage production is low to moderate (Hansen and others 1988a,b). Coyote willow communities often act as firebreaks due to their occurrence on sites with high water tables or near streams, but they may burn during unusually dry years. Burned plants may resprout from roots and spread from offsite seeds dispersed to wet mineral soils (Conrad 1987; Rowe and Scotter 1973; Zasada 1986).

Varieties and Ecotypes—“Silvar” coyote willow Figure 47—Geyer willow forms clumps or corridors was cooperatively released by the USDA Soil that provide cover and travel routes for big game.

676 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families are brownish purple and glabrous. Early leaves of the 1988; Gaffney 1941; Padgett and others 1989). It is season are glabrous above with long, silky hairs be- more palatable than Drummond willow, Wolf willow, neath. Mature leaves are lanceolate to elliptical, en- and Booth willow (Boggs and others 1990). Geyer tire to inconspicuously toothed, 1.2 to 3 inches (3 to willow clumps and corridors provide excellent cover 8 cm) long, and 0.3 to 0.5 inch (8 to 12 mm) wide. and travel routes for big game. Beaver use it for food Surfaces are silvery to gray green above, glaucous and building material (Allen 1983). It also provides below, and hairy on both sides. Catkins expand with food and cover for birds and small mammals (Argus the leaves on short, leafy branchlets. Staminate cat- 1957). Roots of Geyer willow growing along streams kins are yellow to reddish and 0.3 to 0.6 inch (0.7 to 1.5 contribute to formation of overhanging banks; the cm) long. Pistillate catkins are globose, reddish, 0.4 to canopy provides shade for fish. 0.6 inch (1 to 1.5 cm) long, and loosely flowered. The In eastern Oregon palatability of Geyer willow to persistent floral bracts are sparsely hairy and yellow livestock was rated as moderately high (Kovalchik to brown or blackish. Capsules are hairy, ovoid oblong, and others 1988). Willows and understory species in and 0.1 to 0.2 inch (3 to 6 mm) long. Stigmas are nearly these communities are often highly productive and entire (Brunsfeld and Johnson 1985; Hitchcock and may receive heavy use (Hansen and others 1988). others 1964). Geyer willow communities provide easy access to livestock. Early season use by wild ungulates, livestock, or Ecological Relationships and Distribution— vehicles may adversely impact wet organic soils (Man- Geyer willow occurs at low to upper elevations from ning and Padgett 1995). Excessive use may lead to British Columbia to Montana and south from California decreased willow vigor or stand loss, replacement of to Colorado. It grows in wet meadows and marshes, understory graminoids with less desirable species, beaver ponds, valley bottoms along meandering lowering of the water table, erosion, and conversion to streams, and on dry stream benches from the sage- drier community types (Kovalchik 1987). Decadent brush to the spruce-fir zone (Brunsfeld and Johnson Geyer willow can recover if released from browsing 1985; Uchytil 1991b). It most commonly grows on fine- pressure. textured silty to clay-loam soils that may contain some Plants recover from burning or mechanical damage cobbles or gravels and have considerable organic ma- by sprouting or possibly by establishment of seedlings terial and mottling near the surface (Padgett and from offsite seed sources (Kovalchik and others 1988). others 1989; Youngblood and others 1985). Plants Fires are infrequent in these areas, but may occur in usually grow in clumps, but may form a more or less dry years. Prescribed burning may be used to rejuve- continuous corridor near streamside. Communities in nate decadent stands. Areas must be protected from broad valleys are relatively stable and maintained by browsing until the stands recover (Boggs and others seasonal flooding and high water tables within 3.3 ft 1990; Kovalchik 1987). (1 m) of the surface. Associated willow species often include Bebb willow, Booth willow, yellow willow, Varieties and Ecotypes—None. plainleaf willow, Drummond willow, and Lemmon willow. The understory often includes a moderate to dense cover of graminoids and forbs. Family Salicaceae ______Geyer willow flowers from May to August (Dittberner Salix glauca and Olson 1983; Munz 1973). In eastern Oregon seed Grayleaf willow, glaucous willow dispersal begins in early July (Padgett 1981). Seed- lings colonize wet, exposed, well-aerated mineral soils, Description—Grayleaf willow is a low shrub 1 to 4 but can withstand anaerobic conditions (Manning and ft (0.3 to 1.2 m) tall. Young twigs are grayish and hairy, Padgett 1995). becoming reddish brown and glabrous as they mature. Plant Culture—Either unrooted or rooted Geyer Bark of older plants may be rough and furrowed. cuttings may be used to revegetate disturbed sites. Mature leaves are lanceolate to elliptical, entire to Cuttings form roots in about 10 days and shoots in minutely serrulate, and up to 2.2 inches (5.5 cm) long about 10 to 15 days (Platts and others 1987). Conroy and and 0.3 to 0.9 inch (0.7 to 2.2 cm) wide. Mature leaf Svejar (1991) and Svejcar and others (1992) found that surfaces are pale green above and glaucous beneath. establishment of Geyer willow was best if unrooted They are grayish hairy when young, becoming gla- cuttings were planted with the base within 12 inches brous as they mature. Catkins expand with the leaves (30 cm) of the midsummer water table. Waterlogged and remain on the plant through summer. Staminate conditions had no effect on growth of the cuttings. catkins are 0.6 to 1.6 inches (1.5 to 4 cm) long. Pistil- late catkins are 0.6 to 2 inches (1.5 to 5 cm) long and Uses and Management—Geyer willow is used by densely flowered. Floral bracts are persistent, hairy, moose and elk, especially in winter (Chadde and Kay and pale brown to black. Capsules are 0.2 to 0.3 inch

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 677 Chapter 23 Shrubs of Other Families

(5 to 7 mm) long, long beaked, and densely hairy, 1980). Grayleaf willow often becomes highly abun- becoming nearly glabrous as they mature. Stigmas are dant following fire, establishing from seed sources in bilobed (Brayshaw 1976; Brunsfeld and Johnson 1985; unburned areas (Foote 1983). Seedlings have been Hitchcock and others 1964; Viereck and Little 1972). noted to invade disturbed sites in Arctic areas In Alaska and Yukon, grayleaf willow flowers in (Densmore and Zasada 1983; Viereck and Little 1972). June. Fruits ripen in July and August and seeds are Grayleaf willow sprouts following mechanical dam- dispersed in late August and September (Densmore age or burning (Haeussler and Coates 1986). It also and Zasada 1983; Viereck and Little 1972). spreads vegetatively through rooting of horizontal stems (Sampson and Jones 1977). Ecological Relationships and Distribution— Grayleaf willow is a circumboreal species, extending Plant Culture—Densmore and Zasada (1978) found south in North America to alpine and subalpine sites that untreated cuttings of 2-year-old wood, taken in in eastern Idaho, Utah, Montana, Wyoming, Colorado, autumn following leaf fall or in spring following leaf and northern New Mexico. In the Intermountain area expansion, produced few roots. Cuttings treated with it occurs along streams and in other wet places, on rooting hormones produced few to moderate numbers of talus slopes, and in snow concentration areas in alpine roots along the entire length of the cutting (Platts and meadows and forest openings (Padgett and others others 1987). Roots and stems appeared in about 10 1989; Welsh and others 1987). It is found on soils days. Cuttings have been used to plant unstable sand ranging from well drained to poorly drained and usu- dunes in northern Alberta (Hardy BBT Limited 1989). ally waterlogged (Alaska Rural Development Council Seed may be used to propagate planting stock. Direct 1977). A low-growing form occurs in open, alpine seeding may also be practical (Uchytil 1992). situations (fig. 48); plants in somewhat protected, Densmore and Zasada (1983) and Zasada and Viereck subalpine environments are more erect and shrublike (1975) found that fresh seeds were conditionally (Dorn 1970). An early successional species, grayleaf dormant. Germination increased with constant incu- willow pioneers fresh alluvial deposits, burns, and bation temperatures from 41 to 77 ∞F (5 to 25 ∞C). Wet other disturbed areas with exposed mineral soils chilling for 90 days released dormancy; germination (Viereck and Little 1972). was initiated during wet chilling, and nearly all seeds Densmore and Zasada (1983) and Zasada and germinated when incubated at constant temperatures Viereck (1975) reported that grayleaf willow pro- in the 41 to 77 ∞F (5 to 25 ∞C) range. Environmental duces large numbers of small, lightweight seeds that conditions may influence the level of seed dormancy; remain on the plant through summer and are dis- this was indicated by variability in germination of persed in fall. These seeds remain viable through seeds by year of collection and geographic origin. winter and germinate in spring following snowmelt; Germination also varied among seeds from different this permits seedling development to occur over the catkins on the same plant. entire growing season. Spring ripening populations, Seeds may be stored at 14 ∞F (–10 ∞C) in doubled 3- however, have been reported (Watson and others mil polyethylene bags for up to 3 years without loss of viability (Zasada and Densmore 1980; Zasada and Densmore 1977). The initial dormancy of some seed lots is lost in storage. Use and Management—Grayleaf willow is browsed by moose, but it is generally less palatable than other co-occurring willows (Milke 1969). Caribou make use of the species, primarily in summer; snowshoe hare use is concentrated in winter (Cody 1965; Smith and others 1988). Nutrient quality of grayleaf willow for ungulates is good in winter (Risenhoover 1987; Scotter 1972), but lower growing forms may be covered by snow. Plants are tolerant of heavy browsing. The cover value of grayleaf willow for large ungulates is also limited by its low-growing habit, but dense stands provide good cover for small animals (Uchytil 1992). Erosion control value of grayleaf willow is considered moderate due to its intermediate growth rate (Alaska Rural Development Council 1977). Estab- Figure 48—Grayleaf willow occurs in alpine areas lished plants, however, provide good soil stabilization. and exhibits a low, spreading growth habit. Varieties and Ecotypes—There are no releases.

678 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Family Salicaceae ______five stamens. Pistillate catkins are pale green to yellow or white and 0.8 to 2.8 inches (2 to 7 cm) long. Salix lasiandra var. caudata Floral bracts are deciduous, yellow green, hairy on the Whiplash willow lower part, and glabrous distally. Capsules are narrowly ovate, glabrous, and 0.16 to 0.3 inch (4 to 8 mm) Description—Salix lasiandra plants growing in long. Stigmas are lobed. Flowering occurs in April or the Intermountain region belong to S. l. var. caudata, May, and fruiting and seed dispersal from June to whiplash willow. Whiplash willow ranges from 10 to August (Brinkman 1974i; Hitchcock and others 1964). 20 ft (3 to 6 m) in height (fig. 49). It is treelike at Pacific willow (S. l. var. lasiandra) occurs mostly lower elevations, becoming shrubby at higher eleva- west of the Cascade Mountains, but also in wetter tions. Young twigs are finely hairy, becoming glabrous portions of eastern Washington, northern Idaho, and and yellowish with age. Large branches are yellow to northwestern Montana. It is distinguished by the red or brown in winter, while small branches are glaucous lower surface of its leaves (Brunsfeld and orangish (Sutton and Johnson 1974). Mature leaves Johnson 1985). are lanceolate to elliptic, widest at or below the middle, and serrulate. Leaf surfaces are green and glabrous. Ecological Relationships and Distribution— The lower surface is paler than the upper, but not Whiplash willow is distributed from Alaska to glaucous. Petioles often bear glands on the upper side Saskatchewan and south from southern California to near the junction with the leaf blade. Catkins expand New Mexico at low to mid elevations (Welsh and with the leaves. Staminate catkins are white to yellow others 1987). Associated uplands support communi- and 0.6 to 1.8 inches (1.5 to 4.5 cm) long, with three to ties ranging from Wyoming big sagebrush to pinyon- juniper, mountain big sagebrush, and Douglas-fir (Uchytil 1989b). Whiplash willow is commonly a pioneer or early seral species on alluvial sands and gravels adjacent to fluctuating streams and rivers or abandoned river channels (Brunsfeld and Johnson 1985). Stands are maintained by periodic flooding. Whiplash willow may maintain its dominance on stabilized sandbars, as they develop into benches within the flood plain, but it is eventually replaced by later seral willow or cottonwood species (Manning and Padgett 1995). It often occurs with coyote willow, another pioneer species; they form intermittent linear communities between the high water level and later successional woody species growing further from water. Whiplash willow exhibits high tolerance to deposition and flooding and low tolerance of drought and salty soils (USDA Soil Conservation Service 1992). Its productivity is generally low due to repeated flooding. Whiplash willow spreads primarily from seeds that are dispersed by wind and water (Zasada 1986). Bro- ken twigs or branches of whiplash willow transported downstream and deposited on wet alluvial surfaces may sprout, developing new plants vegetatively (Argus 1973). Plant Culture—Whiplash willow is widely used for stabilizing streambanks due to its early seral status. It may also be used on appropriate sites to provide naturalized landscaping, screens, or windbreaks. Unrooted or rooted cuttings may be used. Unrooted cuttings are most successful in areas with high water tables throughout the summer and a long growing season. Whiplash willow roots readily along the entire length of the stem. Roots initiate growth in about 10 Figure 49—Pistillate catkins of whiplash willow are days and shoots in 10 to 15 days (Platts and others conspicuous, white to green, and appear with the 1987). leaves.

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Seeds are green when preripe and yellowish when rocky soils where soil water is adequate. Nehalem is ripe (Brinkman 1974i). There are about 11,500,000 tolerant of flooding and may be planted in reservoir cleaned seeds per lb (25,300,000 per kg). Brinkman drawdown areas. Stands may be established from (1974i) reported 25 percent germination at 72 ∞F cuttings; suitable materials may be used for wattling, (22 ∞C) in 3 days. Densmore and Zasada (1983) found brush matting, and branch packing to provide slope that fresh seed collected in Alaska germinated rapidly protection. and nearly completely at constant incubation tem- “Roland” whiplash willow was released in 1985 by peratures ranging from 41 to 77 ∞F (5 to 25 ∞C). the Alaska Plant Materials Center (Wright 1989). It is used for landscaping and stream revegetation and Uses and Management—Whiplash willow colo- protection throughout Alaska. nizes and stabilizes recent alluvial deposits. Roots provide streambank stability; the canopy provides shade for fish. Consequently, management to main- Family Salicaceae ______tain these stands is critical. Whiplash willow provides valuable habitat for deer and nongame birds (Argus Salix lasiolepis 1973; Arno and Hammerly 1977; Bernard and Brown Arroyo willow 1977; Gray and Greaves 1984). Although palatability of whiplash willow for wildlife is generally considered Description—Arroyo willow is a shrub or small poor or fair, it is an important browse for mule deer tree 13 to 20 ft (4 to 6 m) tall (fig. 50) with multiple (Sampson and Jespersen 1963; USDA Forest Service stems, a large root crown, and a fibrous root system. 1937). Beaver use it heavily in winter (Kindschy 1985). Young twigs are hairy and yellowish olive to reddish. Palatability of whiplash willow is greater for sheep Mature leaves are coriaceous, oblong or oblanceolate, than for cattle. On wet sites, healthy whiplash willow and entire or rarely minutely serrate with somewhat stands are not easily accessed by livestock because of revolute margins. They are 0.6 to 1.6 inches (1.5 to 4.1 the dense overstory. Therefore, these stands are only cm) long and 0.2 to 0.5 inch (5 to 13 mm) wide. Leaf moderately susceptible to overgrazing, trampling, surfaces are dark green, glabrous above, and glaucous and compaction (Manning and Padgett 1995). beneath. Catkins expand before to slightly after the Communities with coarse, dry surface soils and lim- leaves. Staminate catkins are 0.9 to 1.8 inches (2.2 to ited herbaceous cover are more vulnerable to heavy 4.5 cm) long. Pistillate catkins are densely flowered grazing. With prolonged heavy use, willow recruit- and 0.9 to 1.8 inches (2.2 to 4.5 cm) long. The persistent ment is decreased, mature plants are highlined, and floral bracts are purplish black and densely hairy. plants become decadent or die. Willows may be re- Capsules are ovate, glabrous, and 0.1 to 0.2 inch (3 to placed by more mesic shrubs, introduced grasses such 5 mm) long (Hitchcock and others 1964; Welsh and as Kentucky bluegrass, or weeds (Hansen and others others 1987). 1995). With reduction or elimination of browsing, In northern Arizona, arroyo willow flowers from existing plants often recover quickly. mid-March to mid-April (Sacchi and Price 1992). Seeds The response of whiplash willow to burning has been are dispersed over a 3-week period from late April to poorly documented, and its resprouting capability is mid-May. not known. Windblown seeds play an important role in reestablishing the species in burned or disturbed areas (Miller and Miller 1976; Shaw and Clary 1996; Zasada 1986). Varieties and Ecotypes—“Nehalem” Pacific willow is a male clone released by the USDA Soil Conser- vation Service Corvallis Plant Materials Center in 1978 (Darris and Lambert 1993). It originated from a native population growing near the Nehalem River in northwestern Oregon. Nehalem was selected for its high basal stem density and attractive foliage. It may be used to stabilize streambanks, improve riparian and aquatic habitat for wildlife and fish, and provide natural area landscaping and screens. Nehalem is adapted to planting sites on sandbars, lakeshores, and riverbanks at elevations below 4,900 ft (1,500 m) on the west side of the Cascade Mountains from Wash- ington to northern California. It may be planted on Figure 50—Arroyo willow grows along washes, ditches, soils ranging from sandy loams to gravelly or rich, and well-drained sites along ephemeral streams.

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Ecological Relationships and Distribution— California on the west side of the Cascade Mountains. Arroyo willow is distributed at low elevations from The release is propagated from cuttings and can be British Columbia to Idaho and south from Baja Cali- used for wattling, brush matting, and branch packing fornia to west Texas and northern Mexico. It occurs in combination with mechanical site treatments (Darris entirely east of the Cascade Mountains. Arroyo willow and Lambert 1993). may be found growing along streams, ditches, and washes (Welsh and others 1987). It is adapted to well- drained, ephemerally wet riparian sites (Manning and Family Salicaceae ______Padgett 1995). In Nevada, arroyo willow occurs on Salix lutea streambenches with coarse-textured or dry surface Yellow willow, shining willow soils, along incised or ephemeral streams, and near seeps (Manning and Padgett 1995). Description—Yellow willow is a rounded shrub or Plant Culture—Arroyo willow may be propagated rarely a small, usually multi-stemmed tree 10 to 20 ft from cuttings. Rooting ability ranges from erratic (3 to 6 m) tall (fig. 51). Young twigs are sparsely hairy (Platts and others 1987) to very good (USDA Soil and yellowish to reddish, becoming glabrous and yel- Conservation Service 1992). Roots form from the cal- lowish white, gray, or brownish with age. Mature lus and over the lower one-third of the stem (Platts and leaves are lanceolate to elliptical, entire to serrulate, others 1987). Stem and root growth are initiated in 0.8 to 2.2 inches (2 to 5.5 cm) long, and 0.4 to 0.8 inch about 10 days. (10 to 21 mm) wide. They are dark green to yellow Seed biology of arroyo willow has received little green above and pale green and glaucous below. How- study. Seeds are nondormant and remain viable in ever, a nonglaucous form has been reported (Manning nature for only 1 to 3 weeks (Sacchi and Price 1992). Uses and Management—Arroyo willow has good habitat value, high tolerance to flooding and sediment deposition, moderate drought tolerance, and low salt tolerance (USDA Soil Conservation Service 1992). Management to preserve arroyo willow communities in otherwise arid areas is critical; they provide important habitat for a wide range of invertebrate and vertebrate species. Native seedlings of arroyo willow establish on fresh fluvial deposits and open or partially vegetated streambanks (Sacchi and Price 1992). High seedling mortality is associated with drying of the soil surface; competition and herbivory play minor roles. Seedling growth is reduced by shading, and small seedlings have low overwinter survival. Seedlings surviving for 3 growing seasons may be considered established. Varieties and Ecotypes—“Rogue” arroyo willow was cooperatively released by the Corvallis USDA- SCS Plant Materials Center in 1990 for use in stabilizing streambanks of low velocity meandering streams, improving freshwater fish and wildlife habitat, naturalized landscaping, and windbreaks or screens in riparian or moist upland situations (Darris and Lam- bert 1993). Rogue is a male clone derived from a population growing along the Rogue River in Curry County, OR (Darris and Lambert 1993). Plants are large, multistemmed or occasionally single trunked with spreading crowns. Rogue was selected for its rapid, early growth rate and low incidence of insect and disease pests. It is widely adapted to riparian areas and moist Figure 51—A low- to mid-elevation species, yellow upland sites receiving greater than 35 inches (890 mm) willow frequently occurs as an early seral species on of annual precipitation from Washington to northern periodically flooded streambars.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 681 Chapter 23 Shrubs of Other Families and Padgett 1995). Catkins expand slightly before or may be shifted to shrubs such as Woods rose, mesic with the leaves. Staminate catkins are 0.8 to 2 inches forbs, or with prolonged excessive grazing, Kentucky (2 to 5 cm) long. Pistillate catkins are 0.8 to 2.8 inches bluegrass and other weedy and exotic species. (2 to 7 cm) long and densely flowered. The persistent Plant Culture—Yellow willow may be planted as floral bracts are hairy and brown to blackish red. unrooted or rooted cuttings. Unrooted cuttings are Capsules are glabrous and 0.1 to 0.2 inch (3 to 6 mm) better able to establish on sites with a high water table long, produced on stipes less than 0.1 inch (2.5 mm) that persists throughout the summer and allows roots long. Stigmas are often scarcely lobed (Brunsfeld and to develop. Roots are initiated along the entire length Johnson 1985; Hitchcock and others 1964; Welsh and of the stem, but are most abundant along the lower others 1987). one-third. Under greenhouse conditions, new roots Flowering and fruiting dates vary within and among and stems are initiated in about 10 days (Platts and populations (Martens and Young 1992). Catkins ap- others 1987). Although cuttings root rapidly, only pear in late March or early April on stands near Reno, moderate numbers of roots are produced. NV. Flowering has been reported to occur from May Seed of yellow willow may be collected by harvest- to June in California (Munz 1973). Capsules ripen ing the inflorescences as the capsules begin to open about a month following flowering (Martens and (Martens and Young 1992). Developing fruits should Young 1992). Catkins on a single branch vary in be monitored carefully because seed is dispersed phenological development, and flowering within each quickly when mature. Harvested catkins can be spread catkin is indeterminate. Consequently, fruit ripen- to dry in a warm room until capsules open, releasing ing is not uniform. Seeds are dispersed by wind and the seed. are buoyant in water. Hairs on the seed may aid in Fresh yellow willow seed harvested near Reno, NV, dispersal, but they may also be important in germi- exhibited greatest germination in the dark at 50 to nation and seedling establishment (Martens and 77 F (10 to 25 C) (Martens and Young 1992). Germi- Young 1992). ∞ ∞ nation at 36 ∞F (2 ∞C) exceeded 10 percent; no seeds Ecological Relationships and Distribution— germinated at 95 to 104 ∞F (35 to 40 ∞C). When Yellow willow is distributed from Washington to germinated in the dark at 55 alternating temperature Manitoba and south from California to Nebraska regimes, utilizing temperature combinations between (Brunsfeld and Johnson 1985). A common species 32 and 104 ∞F (0 and 40 ∞C) (8 hrs/16 hrs), greatest within its range, it grows along streams and ditches in germination (52 percent) occurred at 50/36 ∞F (10/2 ∞C). valleys and canyon bottoms. It generally occurs at low Temperatures corresponding to very cold, cold, and to mid elevations, but may be found at higher eleva- moderate seedbed conditions enhanced germination; tions, particularly in the southern portion of its range. alterations corresponding to warm and warm/fluctu- Associated upland communities include mountain big ating seedbed conditions reduced germination. sagebrush, pinyon-juniper, mountain mahogany, Yellow willow seeds can be stored for short periods Jeffrey pine, or lodgepole pine (Uchytil 1989c). of time (Martens and Young 1992). Seeds exhibited Yellow willow is capable of withstanding prolonged some viability after 7 weeks when stored in water at inundation and often occurs as a pioneer or early seral 36 ∞F (2 ∞C), in paper bags at 32 ∞F (0 ∞C), or at room species on periodically flooded streambars. On such temperature (Martens and Young 1992). Viability of sites it may occur with, or replace, whiplash willow or seeds stored in a desiccator over calcium chloride at coyote willow. Understory cover on such sites is often 32 ∞F (0 ∞C) declined from 25 to 2 percent after 2 weeks. sparse. Yellow willow stands also occur on wet alluvial Uses and Management—The USDA Soil Conser- benches or terraces with well-developed, usually fine- vation Service (1992) described yellow willow as hav- textured soils. Water tables are generally high, but ing good habitat value, good rooting ability, moderate may fall below 3.3 ft (1 m) in summer. Understory tolerance of flooding, moderate drought tolerance, and species include beaked sedge, Nebraska sedge, bluejoint low salt tolerance. Yellow willow often forms a thick reedgrass, and other graminoids (Hansen and others corridor that provides streambank stabilization. The 1995; Manning and Padgett 1995; Youngblood and fibrous root system contributes to the development of others 1985). Understory productivity and accessibil- overhanging banks; the canopy provides shade and ity to livestock depend on the extent of willow canopy. litter, enhancing fish habitat. Yellow willow communities may occur in mosaics with A dense canopy cover of yellow willow provides food other riparian community types dominated by cotton- and cover for big game, beaver, birds, and other wild- woods, Booth willow, or Drummond willow. life. Moose and elk use yellow willow in summer and With excessive livestock grazing in yellow willow winter. Excessive browsing, however, can result in communities, the water table falls, surface soils dry, stand opening or loss of willows (Gaffney 1941; Hansen and streams may become incised (Hansen and others and others 1995; Van Dersal 1938). 1995; Manning and Padgett 1995). Associated species

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Livestock may make heavy summer and fall use of Three varieties are recognized. Salix planifolia var. palatable understory species associated with some planifolia ranges from 6.6 to 13 ft (2 to 4 m) in height yellow willow stands. Heavy livestock use of willows with elongate leaves 1.4 to 3.1 inches (3.5 to 8 cm) long. may reduce their vigor, or if prolonged, result in stand The leaves are usually red tinged and sparsely hairy. loss. Wet soils are easily damaged by livestock tram- The other two varieties are generally less than 6.6 ft pling early in the season, resulting in soil compaction, (2 m) in height. S. p. var. monica is usually 3.3 ft (1 m) early drying of the soil surface, and bank sloughing. or less in height with leaves that are 1 to 1.4 inches (2.5 Decadent stands resulting from excessive browsing by to 3.5 cm) long. S. p. var. pennata is 3.3 to 6.6 ft (1 to livestock or big game will regenerate if browsing 2 m) tall with leaves 1.8 to 2.6 inches (4.5 to 6.5 cm) pressure is reduced. Fire may also be used to rejuve- long (Brunsfeld and Johnson 1985; Hitchcock and nate decadent stands (Hansen and others 1995). Rec- others 1964). reation and travel within these communities require Ecological Relationships and Distribution— careful consideration due to the dense willow canopy Plainleaf willow is circumboreal and extends south- and early season wet soils (Hansen and others 1995). ward in North America from California to New England Campsites and trail or road routes should skirt these (Welsh and others 1987). It is found along streams, areas when possible. around the margins of lakes and ponds, and in other Varieties and Ecotypes—None. wet areas. Salix planifolia var. planifolia occurs east of the mountains in Western Canada and the Northwestern Family Salicaceae ______United States at low to moderate elevations. In the Salix planifolia United States, it is distributed from the upper sage- Plainleaf willow brush to the Douglas-fir zone in northern Idaho, Mon- tana, and Wyoming (Uchytil 1991c). It grows on min- Description—Plainleaf willow is a shrub growing eral soils with textures ranging from gravelly to sandy to 13 ft (4 m) in height. Twigs are glabrous and black or clayey. Water tables on these sites may fall below to purplish black, exfoliating in translucent flakes. 3.3 ft (1 m) by midsummer. Associated species often Mature leaves are elliptical, entire, dark green, shiny include other tall willows such as Bebb willow, Booth above, strongly glaucous beneath, and not perma- willow, or Geyer willow. nently pubescent on both sides (fig. 52). Veins are Salix planifolia var. monica occurs in valley bottoms prominent and partially parallel. Catkins expand beat high elevations and on wet, open, subalpine slopes fore or with the leaves. Staminate catkins are 0.4 to from central Idaho to west-central Montana, and south 1 inch (1 to 2.5 cm) long. Pistillate catkins are densely from the Sierra Nevada Mountains of California to flowered and 0.8 to 1.6 inches (2 to 4 cm) long. Floral New Mexico. It often occurs in upper cirque basins bracts are persistent, black, and long hairy. Capsules with cold air drainages (Brunsfeld and Johnson 1985). are short hairy and 0.1 to 0.3 inch (3 to 7 mm) long. It usually grows on wetter sites than S. p. var. plani- Stigmas are lobed (Hitchcock and others 1964). folia. S. p. var. monica is often associated with Wolf willow, but grows on saturated soils with finer textures. Soils on S. p. var. monica sites usually have an organic surface horizon overlying alluvial sands, silts, clays, or sometimes gravels. Soils are often gleyed near the upper horizon (Youngblood and others 1985). Soil pH is generally slightly to strongly acidic (pH 4.4 to 6.3). Stands may be flooded in spring, and the water table generally remains within the rooting zone throughout the growing season. Lowering the water table may lead to an increase in Wolf willow. S. p. var. pennata is more narrowly distributed. It occurs in the Cascade Mountains from northern Washington to northern Oregon. Plant Culture—Rooted or unrooted plainleaf willow cuttings may be planted. Unrooted cuttings root rapidly, but rooted cuttings are recommended for high-elevation sites with short growing seasons and Figure 52—Plainleaf willow is a small- to medium- rapidly drying soils. Stems produce low to moderate sized shrub with dark green elliptical leaves that are numbers of roots along their entire length (Platts and shiny above and glaucous beneath.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 683 Chapter 23 Shrubs of Other Families others 1987). Root formation is initiated in about 10 Protection from grazing by livestock during the grow- days; shoots require 10 to 15 days to develop under ing season leaves a thick cover of dry vegetation that greenhouse conditions. burns well in spring before new growth is initiated. Densmore and Zasada (1983) found that seeds col- Burns along streambanks should be conducted with lected in Alaska were nondormant at maturity. Ger- caution to avoid loss of the bank stabilization provided mination was rapid and nearly complete at constant by these communities. Burned sites should be pro- temperatures ranging from 41 to 77 ∞F (5 to 25 ∞C). tected from grazing for 2 to 3 years to prevent overuse of recovering plants. Uses and Management—Roots of plainleaf willows growing in riparian areas contribute to the for- Varieties and Ecotypes—None. mation of overhanging banks. Plants also furnish shade for fish habitat. Beaver use the plant for food and building materials. Stem pieces not consumed Family Salicaceae ______may sprout and develop into new plants (Cottrell Salix scouleriana 1995). Big game use plainleaf willow communities for Scouler willow, fire willow, black willow, forage, cover, and as travel corridors (Hansen and others 1995; Manning and Padgett 1995; Padgett and mountain willow, nuttall willow others 1989). Moose make heavy winter use of S. p. Description—Scouler willow is a shrub or var. planifolia, but use by elk and mule deer is low multistemmed tree arising from a massive root crown (Dorn 1970; Mattson 1984; Van Dersal 1938). Winter and a deep, spreading root system. Mature trees may use of S. p. var. monica may be limited due to its low be 5 to 60 ft (1.5 to 18 m) tall. Young twigs are gray and stature. However, in some areas, moose browse all short hairy, becoming dark reddish brown and gla- branches that are accessible in winter. Plainleaf wil- brous. The crushed bark has a distinctive skunklike low communities also provide valuable cover and food odor. Mature leaves are oblanceolate to lanceolate for small mammals and birds (Douglas and Ratti 1984; or ovate, entire to crenate or serrate, 0.8 to 2.4 Finch 1987). inches (2 to 6 cm) long, and 0.4 to 1.2 inches (1 to 3 cm) Open stands of S. p. var. planifolia provide moder- wide. Leaf surfaces are dark green and glabrous above ate amounts of forage for livestock (Hansen and others and sparsely reddish hairy beneath. Leaves turn yel- 1995) compared to other willow communities. Produc- low in autumn. Catkins usually expand before the tion of these communities is moderate. Palatability of leaves and are soon deciduous. They are nearly sessile. S. p. var. planifolia to cattle is low (Dorn 1970), but Staminate catkins are yellowish white, 0.6 to 1.4 palatability of some associated understory species is inches (1.5 to 3.5 cm) long, and about as long as wide. high. Early season or season-long use by livestock Pistillate catkins are reddish, densely flowered, 0.8 to should be avoided as trampling of saturated sites will 2.4 inches (2 to 6 cm) long, and 0.5 to 0.7 inch (1.3 to 1.7 damage plants and soils (Manning and Padgett 1995). cm) wide. Floral bracts are brown to black and long Short growing seasons and continually wet soils usu- hairy. Capsules are hairy, 0.2 to 0.3 inch (5 to 8 mm) ally limit livestock use to late summer and early fall long, and more or less long beaked. Stigmas are entire (Hansen and others 1995). The grazing value of sites to bilobed (Brunsfeld and Johnson 1985; Hitchcock dominated by S. p. var. monica is limited due to its and others 1964). small stature, low productivity, and association with Scouler willow is one of the earliest willows to flower. wet soils. Flowers appear from April to July (Orme and Leege Subalpine and alpine areas occupied by plainleaf wil- 1980); fruits ripen and seeds disperse from May to July low provide an array of recreational opportunities in- (Brinkman 1974i). Seeds remain viable for only about cluding hiking, wildlife viewing, and fishing (Hansen 1 week (Densmore and Zasada 1983). and others 1995). These communities, particularly the wetter S. p. var. monica sites, are easily damaged by Ecological Relationships and Distribution— hikers, livestock, horses, and offroad vehicles. Heavy use Scouler willow is distributed from Alaska to Manitoba can cause soil compaction, rutting, loss of vegetation, and south to California, South Dakota, and New Mexico formation of multiple trails or roads, streambank ero- (Brunsfeld and Johnson 1985). At low to mid eleva- sion, and other long-term damage that cannot be easily tions, it occurs in valley bottoms, along streams, and repaired. Travel should be restricted to existing trails or around springs, where it attains a treelike stature. roads, and these carefully maintained. New travel routes Scouler willow, however, is generally an upland spe- should be constructed on adjacent uplands. cies, growing on well-drained slopes at higher eleva- Response of the species and its varieties to burning tions, where it occurs as a minor understory species in and its ability to resprout have not been examined late seral Douglas-fir, subalpine fir, western hemlock, (Hansen and others 1995). Short-term productivity of and western red cedar forests (Arno and Hammerly understory sedges can be increased by burning. 1977; Froiland 1962). It is slightly tolerant of shade

684 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families and capable of persisting on these sites at low frequen- growing seasons (Leege and Hickey 1971; Lyon 1971). cies (Cooper and others 1987; Steele and Geier-Hayes Vegetative spread occasionally occurs through rooting 1989, 1992). Scouler willow may be found on mineral of branch segments (Watson and others 1980). soils of varying textures and on peaty soils (fig. 53). It Dense stands of Scouler willow seedlings and regen- is tolerant of acidic, but not salty, soil conditions. erating plants may form shrub fields following fire. Scouler willow proliferates following disturbance. It These may persist for varying periods of time with often becomes widespread following stand-destroying willow coverage decreasing as conifers mature (Steele wildfires, slash burning, clearcutting, road building, and Geier-Hayes 1995). or other disturbances that expose mineral soil and Plant Culture—Scouler willow has been used to provide suitable sites for seed germination and estab- stabilize cut-and-fill areas along logging roads and lishment of seedlings (Arno and others 1985; Steele other disturbed upland areas, but cuttings do not root and Geier-Hayes 1989, 1992, 1995). It is particularly as readily as those of many other willows (Monsen abundant where soils have been disturbed in such a 1975; Plummer 1976). Roots develop in moderate manner as to trap water. Some broadcast burn opera- numbers from the callus area; they do not develop tions and wildfires that leave 50 percent or more of along the length of the stem. Formation of roots nor- the preburn overstory intact, however, do not provide mally requires about 10 to 15 days under greenhouse suitable seedbeds for Scouler willow, and an increase conditions (Platts and others 1987). Densmore (1978) in snowbrush ceanothus may be favored (Forsythe found that total percent and rate of rooting were 1975; Steele and Geier-Hayes 1995). greater for cuttings of second-year wood harvested in High seedling densities on exposed, wet mineral the spring following leaf expansion (10 percent in 30 soils may result from heavy seed rain originating from days, 1 root each) than for dormant cuttings harvested unburned areas (Lyon 1971; Stickney 1986). Scouler in the fall (4 percent in 60 days, 4.5 roots each). Second- willow seeds sown on burns of differing severities on year wood was collected from a population near upland black spruce sites in Alaska germinated only Fairbanks, AK. on moderately (organic soil layers partially consumed) In a field test, 8- to 10-inch (20- to 25- cm) dormant and severely (ash layer present, organic material in cuttings of 2- to 3-year-old-wood harvested near soil consumed or nearly so to mineral soil) burned Fairbanks, AK, were stored fully imbibed at 14 F (–10 C) seedbeds (Gruell and others 1982; Lyon and Stickney ∞ ∞ until planted soon after soil thaw. Cuttings were 1976). Germination was greatest and seedlings sur- planted vertically with 0.8 to 2 inches (2 to 5 cm) of vived after 3 years only on severely burned sites. stem above ground. Survival following one growing Burned Scouler willow plants resprout vigorously season was 17 percent. Nearly 100 percent of all stem from the rootcrown; multiple sprouts sometimes ap- cuttings collected in late fall and early winter from pear a few days after a fire (Foote 1983). Some plants sites in Idaho stored in a moist cold environment, and regenerate even following burns that destroy the en- spring planted in nursery beds, rooted and produced tire canopy (Lyon and Stickney 1976). Regenerating healthy plants. Plants grown under nursery condi- trees grow rapidly, particularly during the first two tions for one season developed large, robust root systems. When outplanted, the rooted plants established quickly and provided cover on road cuts and fill slopes. Scouler willow may also be propagated from seed. There are about 6,500,000 seeds per lb (14,300,000 per kg) (Brinkman 1974i). Germination of seeds incubated at 85/ 70 ∞F (29/21 ∞C) reached 95 percent in 1 day. Densmore and Zasada (1983) obtained rapid and nearly complete germination of an Alaskan collection when seeds were incubated over a range of constant temperatures from 44 to 71 ∞F (5 to 25 ∞C). Uses and Management—Scouler willow is a highly valued browse species for big game. Seral brush fields of Scouler willow and other shrubs are important winter ranges (Leege 1968). Moose, elk, white-tailed deer, and mule deer use the species throughout the year, but primarily in winter (Kufeld 1973; Kufeld and others 1973; Singer 1979; Smith Figure 53—Generally an upland species, Scouler 1953). In south-central Alaska, moose sometimes willow grows on dry slopes as an understory in remove the bark (Viereck and Little 1972). Recent conifer forests. sprouts are most palatable; greatest production may

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 685 Chapter 23 Shrubs of Other Families occur on branches browsed the previous year. Much when mature. Mature leaves are entire, elliptical to of the browse on mature plants is unavailable be- lanceolate or oblanceolate, 0.5 to 1.6 inches (1.2 to cause plants are quite tall. Scouler willow also pro- 4.2 cm) long, and 0.2 to 0.5 inch (0.5 to 1.3 cm) wide. vides cover for birds and small mammals (Steele and They are gray green to silvery and sparsely to densely Geier-Hayes 1992; Vories and Sims 1977). In seral hairy on both sides. Catkins expand with the leaves. brush fields Scouler willow provides excellent cover Staminate catkins are yellowish and 0.2 to 0.6 inch for big game. Isolated plants in late-seral coniferous (0.5 to 1.5 cm) long. Pistillate catkins are globose, stands are not as effective. Scouler willow is highly fuzzy, white, densely flowered, and 0.4 to 0.8 inch (1 to palatable to cattle and sheep (Dayton 1931). It may 2 cm) long. Floral bracts are persistent, long hairy, and receive greater use than other willow species due to blackish or pale at the base. Capsules are 0.1 to 0.2 the high accessibility of the willow clumps. inch (3 to 5 mm) long and glabrous or rarely hairy. Prescribed burning in fall or spring has been used to Stigmas are lobed (Hitchcock and others 1964; Sutton stimulate sprouting of Scouler willow shrub fields that and Johnson 1974; Welsh and others 1987). have grown beyond the reach of big game (Leege 1968, Ecological Relationships and Distribution— 1969, 1979a). Spring burning may be preferable; fall Wolf willow is distributed from northeastern Oregon burning destroys winter browse. Considerable growth south to Nevada and east to Montana and Colorado may occur during the first growing season following (Brunsfeld and Johnson 1985). In central Idaho it spring burning. Leege (1979a) found that sprouts 5 ft occurs from cooler sites at mid elevations in the Dou- (1.5 m) long were produced following a spring burn in glas-fir zone to subalpine areas, where it grows along Idaho. Burning at 5-year intervals did not decrease streams and around lakes and ponds. Wolf willow productivity (Leege 1968). Plant nutritive value in- community types in southeastern Idaho, northern creases only slightly after burning, but new shoots are Utah, western Wyoming, and Montana occur in broad within reach and more palatable than older shoots. meadows, on alluvial terraces, around seeps, and in Scouler willow in clearcuts may enhance shade- old beaver ponds at mid to upper elevations (Hansen tolerant spruce and Douglas fir establishment by pro- and others 1995; Padgett and others 1989; Youngblood tecting plants from wind and temperature extremes and others 1985). Wolf willow often dominates a low (Steele and Geier-Hayes 1989, 1992). In the absence of shrub overstory, although scattered taller willows a conifer overstory, Scouler willow’s growth habit is may be present. At high elevations, Wolf willow may altered from a narrow, upright form to a more shrublike occur in pure stands (fig. 54). Wolf willow occurs on form with a broad, rounded canopy. These vigorous soils with a relatively thick organic surface horizon plants compete strongly with shade-intolerant seed- that develops from deposition of leaves and twigs. lings of ponderosa pine. Because the two species have Some sites with lower amounts of organic matter have similar growth rates for the first 6 to 8 years, the pine fine mineral soil with high water-holding capacity. seedlings may not survive to overtop the willow. Me- Oxygen is provided by lateral movement of water chanical treatments to remove Scouler willow are through such soils. The water table is generally within generally impractical due to the plants’ deep, spreading root system. Consequently, plantations of Douglas- fir are more likely to be successful. Scouler willow has been recommended for use in watershed plantings (Monsen 1975). Rooted cuttings establish quickly when planted on road fills and other disturbances. They grow quickly and provide excellent ground cover on exposed unstable slopes. They also may be used in naturalized landscaping to provide screens or background plantings. Varieties and Ecotypes—None.

Family Salicaceae ______Salix wolfii Wolf willow Description—Wolf willow is a low-statured shrub 2 to 5 ft (0.6 to 1.5 m) tall with a fibrous, spreading root Figure 54—Wolf willow forms dense patches of low- system. Young twigs are yellow to orange and more or statured shrubs in narrow valley bottoms at high less persistently thin hairy, becoming chestnut brown elevations.

686 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

3.3 ft (1 m) of the surface throughout the growing and Olsen 1983). Clusters of arching stems develop on season. Stream channels in Wolf willow communities older specimens. Young branches are divaricate and are often small and meandering. Streambanks are glabrous to pubescent with red to chestnut brown stable due to the high density of willow roots in the soil. bark. Conspicuous transverse cracks develop in the Overland flow is common during snowmelt. bark during the second season; the bark later turns gray and eventually exfoliates. The light green leaves Plant Culture—Wolf willow may be used to reveg- are opposite, sessile, or short petiolate and ovate to etate streambanks and ponds, although growth may oblong with acute to acuminate tips. They are 1 to 3 be slow. Rooted cuttings are advised for use on high- inches (2.5 to 7.6 cm) long and 0.4 to 1.6 inches (1.0 to elevation sites with short growing seasons. Hansen 4.1 cm) wide. The thin blades are entire, 3 to 5 veined, and others (1995) recommended use of dormant 2- to usually glabrous above, but pubescent around the 4-year-old wood. Cuttings should be 12 to 20 inches (30 edges and along veins beneath. to 50 cm) long and 0.4 inch (1 cm) or more in diameter. Attractive terminal cymes of attractive white flow- Rooting is erratic; few to a moderate number of roots ers develop on lateral branches, each with 3 to 11 normally form along the length of the stem (Platts and perfect, regular flowers. Flowers arise from the axils of others 1987). Ten to 15 days are required for root and gradually reduced leaves. There are four white petals, stem initiation. The seed biology of Wolf willow has not 20 to 60 unequal stamens with conspicuous yellow been studied. anthers, and three to five persistent styles that are Uses and Management—Wolf willow communi- more or less connate. Fruit is a woody capsule with ties should be managed to maintain willows along four cells attached to the persistent calyx. The light stream margins and provide vegetative cover through- brown fusiform seeds are numerous (Hitchcock and out the community. Wolf willow communities provide others 1961; Welsh and others 1987). Flowering oc- valuable streambank stabilization (Hansen and oth- curs from late May to July. Fruits develop from July to ers 1995). The low overstory and diversity of plant September or October when the capsules dehisce and species often present in these communities provide seeds are wind and gravity dispersed (Orme and Leege cover for birds, small mammals, and other small 1980; Young and Young 1986). vertebrates and invertebrates. Cover value for big Lewis mockorange exhibits a high degree of local game is fair. Biomass production of Wolf willow is variation in floral and vegetative characteristics. This moderate. The species is palatable to sheep, beaver, variability has been assigned taxonomic status by and big game. Winter use is minimal when plants are some workers (Hitchcock and others 1961). leafless and snow covered. Understory graminoids and forbs vary in palatability. Species such as water Ecological Relationships and Distribution— sedge or tufted hairgrass may receive heavy use by There are about 50 species of Philadelphus in North cattle, but accessibility may be restricted by the dense America, Mexico, Asia, and central Europe. Only two shrub overstory. Excessive grazing may lead to species, Lewis mockorange and littleleaf mockorange replacement of the understory by less palatable spe- (Philadelphus microphyllus), occur in the Intermoun- cies. Grazing and equipment use should be restricted tain region. Lewis mockorange is distributed from until soils are dry as they are easily compacted and become erodible if the vegetative cover is lost. The response of Wolf willow to fire has not been documented (Hansen and others 1995). Varieties and Ecotypes—None.

Family Saxifragaceae ______Philadelphus lewisii Lewis mockorange Description—Lewis mockorange (fig. 55), known as syringa, Lewis syringa, mockorange, or Indian arrowwood was named for Captain Meriwether Lewis, who first collected it on the Clark Fork River near Missoula, MT, on July 4, 1806. Plants are long-lived perennials, ranging from open to densely branched, erect to rounded, fibrous rooted shrubs 3 to 12 ft (0.9 Figure 55—Lewis mockorange survives on a low to 3.7 m) in height (Carson and Peek 1987; Dittberner foothill site following many years of excessive grazing.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 687 Chapter 23 Shrubs of Other Families

British Columbia south to northern California and and Sherlock 1984). Digging rooted suckers and divid- east to Montana (Hitchcock and others 1961; ing the crown of mature plants with a sharp spade or Lackschewitz 1986). It grows from near sea level to axe are alternate means of vegetative propagation. 7,000 ft (2,100 m), but is most common at mideleva- Lewis mockorange is a valuable plant for trans- tions. It is a dominant shrub in some climax ponderosa planting on disturbed steep, rocky, unstable slopes pine communities and a common shrub in some seral where it provides soil stabilization and vegetative ponderosa pine communities (Wright 1978). It is also cover. Seedlings or larger stock are recommended for found in sagebrush deserts, lodgepole pine forests, such sites. Lewis mockorange is also useful for plant- and coastal Douglas-fir and redwood forests. It com- ing in transitional areas of degraded riparian zones. monly occurs in foothills and low mountains on all Uses and Management—Lewis mockorange is not aspects, but primarily on northern and eastern slopes normally grazed heavily by livestock, but in some (USDA Forest Service 1937). The species grows on areas it does receive fair amounts of use by cattle and soils ranging from dry, rocky, gravelly loams on open sheep (Leege 1968; USDA Forest Service 1937). It hillsides, to deep, rich alluvial humic loams near frequently occurs with other species that are more riparian zones (USDA Forest Service 1937). It gener- palatable to big game and, consequently, it may re- ally occurs in small scattered clumps in moist, open, or ceive little use, except under severe conditions. How- partially shaded transitional areas along gullies, can- ever, in some areas it does provide good browse for deer yon bottoms, riparian areas, and seeps (Carson and and elk, especially on winter ranges (Kufeld 1973; Peek 1987; Hopkins and Kovalchik 1983). It also Leege 1968; Marchant and Sherlock 1984; grows on cliffs, talus slopes, and rocky hillsides with Stubbendieck and others 1986). New growth is gener- other shrubs such as serviceberry and ocean-spray ally highly palatable to big game (Leege 1968). Leege (Carson and Peek 1987; Franklin and Dyrness 1973). (1969) reported that elk use of Lewis mockorange Plant Culture—Seeds may be collected by hand increased by 30 percent following a spring burn. The stripping the capsules from the plants after maturity, plants provide cover for birds and other small animals. but before the valves begin to open. Dried capsules are Lewis mockorange is classified as a fire resistant or crushed to release the seeds, and trash is removed survivor species (Fischer and Bradley 1987). Although with an aspirator or fanning mill. There are 3,500,000 shoots may be consumed by fire, Lewis mockorange is to 8,000,000 seeds per lb (7,716,000 to 17,637,000 per capable of resprouting from the root crown or caudex kg) of clean seed (Stickney 1974b). Seed fill is often during the first season following burning (Fischer and low. Seeds may be stored in airtight containers for Bradley 1987; Rowe 1983). Consequently, it is found in periods of up to 1 year. They require an 8-week wet early to late seral as well as in climax communities. prechilling at 41 ∞F (5.0 ∞C) to release embryo dor- Lewis mockorange is not as prolific a sprouter as co- mancy (Stickney 1974b). Germination is tested at 72 to occurring shrubs (Leege 1969). Leege and Hickey 79 ∞F (22 to 26 ∞C). (1971) found that spring-burned Lewis mockorange in Seeds may be broadcast seeded on a rough seedbed northern Idaho seral brush fields (grand fir/pachistima and covered lightly or spot seeded in selected, pre- habitat type) resprouted within 1 or 2 months, while pared areas. They may also be surface seeded using a fall-burned shrubs did not resprout until the following Brillion seeder or similar device. Best results are spring. Sprouts were more numerous following fall obtained if seeds are planted in well-drained sites free burning. The historic burn interval in climax ponde- of herbaceous competition. Seeds may be mixed with rosa pine communities is thought to be 6 to 22 years; other shrub seeds that require shallow or surface thus, Wright (1978) prescribed burns at 10 to 15 year planting. intervals to enhance shrub growth. Bareroot stock may be produced by fall seeding or by Lewis mockorange is the state flower of Idaho and is seeding wet prechilled seeds in spring. The tiny seeds one of the more extensively used native shrubs for may be diluted by mixing them with rice hulls to landscaping (Marchant and Sherlock 1984). It is valued improve uniformity of seeding. Seeds should be cov- for its showy white flowers; the fruits and leaves are less ered very lightly. Seedlings develop rapidly and can be attractive. Dense branches and foliage make it suitable transplanted as 1-year-old stock. Container stock may for hedges. Native Americans used the long woody also be grown from seeds. shoots for arrow shafts (USDA Forest Service 1937). Due to a lack of commercial seed sources and ease of Lewis mockorange seedlings establish well, but vegetative propagation, planting stock is frequently should be protected from competition and browsing. grown from cuttings. Dormant 3 to 4 inch (7.6 to Landscape plants should be pruned after flowering 10.2 cm) hardwood cuttings may be planted in a closed because flowers are produced on twigs of the previous cold frame with bottom heat for rooting. Softwood year. The species is normally free of insect and disease cuttings gathered in early summer or early fall are problems, but seedlings are sometimes susceptible to also easily rooted in a cold frame (Doran 1957; Marchant damping off.

688 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Varieties and Ecotypes—The Colfax germplasm turn reddish with age. The edible fruit is a globose, of Lewis mockorange originated from the Colfax, WA, glabrous, many-seeded berry. Fruit color varies from area. St. Maries germplasm originated from St. Maries, yellow to red orange or black. The minute embryo is ID. Several commercial cultivars of Lewis mockorange embedded in a large amount of endosperm (Goodrich have been developed. “Waterton,” selected from the and Neese 1986; Hitchcock and others 1961; Munz and Waterton Lakes area of Alberta, is a hardy, bushy Keck 1959; Pfister 1974; Welsh and others 1987). shrub with flowers scattered over the crown of the Ecological Relationships and Distribution— plant (Marchant and Sherlock 1984). Philadelphus Golden currant is distributed from north-central coronarius, introduced from Europe, is the commonly Washington to Saskatchewan and South Dakota and cultivated mockorange (Welsh 1987). south to California and New Mexico (Welsh and others Littleleaf mockorange is a smaller, short-lived 1987). Hitchcock and Cronquist (1973) reported that shrub, growing from 3 to 7 ft (0.9 to 2.1 m) in height and the shrub occurs along streambanks and washes, and 4 ft (1.2 m) in width. Shrubs have ascending stems and from grasslands and big sagebrush deserts to ponde- a rounded crown with reddish-tan exfoliating bark. rosa pine forests throughout the Pacific Northwest. In Leaves are narrower than those of Lewis mockorange, California it grows along moist streambanks and in and flowers are solitary or in clusters of three. Littleleaf bottom lands at elevations between 2,500 and 7,800 ft mockorange occurs from Utah to Wyoming and south (760 and 2,400 m) (Munz and Keck 1959). Golden to Texas in pinyon-juniper, mountain brush, aspen, currant occurs in many Utah riparian and palustrine lodgepole pine, and Douglas-fir/white fir forests habitats in greasewood-shadscale, sagebrush, pinyon- (Goodrich 1985; Sutton and Johnson 1974; Welsh and juniper, mountain brush, ponderosa pine, and Douglas- others 1987). Littleleaf mockorange receives some use fir communities at 4,395 to 8,490 ft (1,340 to 2,600 m) by mule deer (Patton and Ertl 1982) and other wildlife. (Goodrich and Neese 1986; Welsh and others 1987). It Littleleaf mockorange may be propagated from seed or occupies similar areas in Colorado and New Mexico, cuttings. It is easily transplanted and is a strikingly and is common in the plains and foothills of South attractive ornamental (Sutton and Johnson 1974). Dakota and Saskatchewan (Harrington 1964). Golden currant is generally not widely abundant Family Saxifragaceae ______(Wasser 1982); it occurs as scattered plants, patches, clumps, and in corridors along waterways. It does not Ribes aureum occur as even a minor component of aspen or conifer Golden currant forests in the Intermountain area (Mueggler 1988; Steele and Geier-Hayes 1987; Youngblood and Mauk Description—There are more than 100 species of 1985), but it does grow in aspen/chokecherry commu- Ribes, primarily occurring in the temperate and colder nities. It normally grows on fertile, well-drained sites regions of the northern hemisphere and the Andes such as moist streambanks, washes, ditches, seeps, mountains of South America. They are particularly and springs. It has been widely cultivated as an common in the Western United States (Pfister 1974; Welsh and others 1987). Species with jointed pedicels, several flowers per raceme, and glabrous to glandular berries that are generally unarmed are grouped as currants. The remaining species, usually armed with spines, are referred to as gooseberries (Hitchcock and others 1961). Golden currant is an unarmed, irregularly shaped, multiple-stemmed shrub 3 to 10 ft (0.9 to 3.0 m) tall (Welsh and others 1987) that spreads by suckering (fig. 56). The reddish bark of young twigs turns gray with age. Leaves are alternate, deciduous, and petiolate. Blades are ovate with a cuneate to chordate base, usually glabrous, and palmately three-lobed with the lobes entire, toothed, or lobed. Ascending bracteate racemes occur on the ends of lateral spur branches with two to 15 flowers (Harrington 1964). Flowers are complete and regular with a spicy odor (Stark 1966). The hypanthum is cylindrical with five spreading calyx lobes alternating with five shorter, Figure 56—Golden currant planted as a low- erect petal lobes. Young flowers are golden yellow, but maintenance species in a recreation site.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 689 Chapter 23 Shrubs of Other Families ornamental (Goodrich 1985; Pfister 1974), and now The degree of dormancy varies among seedlots and occupies drier sites within abandoned farmlands, among seeds within a seedlot; this provides an adap- ranches, and wildland areas. tive advantage. However, this characteristic reduces Although considered a species for moist sites the effectiveness of seed pretreatments. Most seeds (Thornburg 1982), golden currant is found on well- require a long period of wet prechilling to break em- drained soils receiving in excess of 16 inches (40 cm) of bryo dormancy. Germination can be hastened and annual precipitation. It usually occurs on sandy to increased by wet prechilling at low temperatures. A silty loam soils (Thornburg 1982), but is also adapted wet prechilling of 60 to 90 days at 28 to 36 ∞F (–2.2 to to clay-loam soils. Plants generally grow in full sun to 2.2 ∞C) is often used to release embryo dormancy partial shade on soils ranging from slightly acidic to (Pfister 1974). Stidham and others (1982) recom- slightly basic (Dittberner and Olsen 1983; Wasser mended that seeds be prechilled on blotters moistened 1982). with potassium nitrate. Seeds should be fall planted to provide a cold, moist Plant Culture—Golden currant flowers appear period for wet prechilling. Spring plantings should use with or slightly before the leaves in April or May. only prechilled seeds. Seeds are small, round, and Fruits ripen in June or July and are dispersed by birds easily dispensed through most seeders. Golden cur- and mammals (USDA Forest Service 1948). In central rant can be seeded alone or in mixtures with selected Utah berries may ripen as late as mid-August (Plummer shrubs and forbs. It should not be seeded directly with and others 1968). grasses or broadleaf herbs that germinate and grow Fruits or berries are hand harvested by stripping rapidly. Seeds are usually sown at a rate of 0.25 to them from branches or by flailing them into containers 2 lb per acre (0.3 to 2.2 kg per ha), depending on the as soon as they ripen; this reduces losses to birds method of seeding and row spacings. Wildland (Pfister 1974). Few wildland stands are large enough seedings are usually conducted using seed-dribblers to produce many pounds of seed; consequently, culti- mounted on tractors. Seeds are often hand planted vated ecotypes are usually the sources sold for wild- in selected areas. Only a small portion of an area is land plantings. Native stands are capable of producing actually planted using these techniques. good seed crops, but only about 4 lb (1.8 kg) of clean Under nursery conditions, seeds are usually sown at seed are extracted from 100 lb (45.4 kg) of fruits a rate of 60 to 80 seeds per ft2 (650 to 860 per m2) or (Pfister 1974). Seed collection costs are high, and tend 40 viable seeds per linear ft (130 per linear m) of to restrict use of the shrub in large-scale seeding row (Pfister 1974). Seeds should be planted 0.25 inch projects. However, viable seed crops are usually pro- (6 mm) deep on a firm seedbed. Planting on sites duced each year at most locations, and local ecotypes having some surface mulch or debris aids in germina- can be collected and propagated for specific plantings. tion and seedling establishment. Adding mulch to the Most seeds are harvested from cultivated plantings, soil surface is recommended for nursery beds subject fence-line plantings, or irrigated seed fields. Plants in to rapid drying and crusting. these situations generally produce more consistent Seed germination is usually only moderate or fair. and abundant crops. In addition, the bushes are more However, seeds that germinate often do so uniformly, uniform in stature, and are closely spaced, which aids and initial emergence is usually very good even under in harvesting. range or wildland conditions. Compared with most Collected fruits should not be allowed to overheat shrubs, seedlings of golden currant are very persis- prior to extraction. Fruits are processed by maceration tent. Seedlings grow rapidly and generally attain using a Dybvig cleaner and water to separate the berry heights in excess of 6 to 12 inches (15 to 30 cm) the first or pulp from the seeds (Plummer and others 1968). growing season. Although natural thinning occurs, Dried fruit should be soaked in water to aid cleaning seedlings are vigorous; normally enough survive to or maceration. Pulp and empty seeds may be sepa- provide a full stand. Seedlings of few other shrubs are rated from sound seeds by flotation. There are 200,000 as vigorous as this species. to 285,000 seeds per lb (441,000 to 628,000 per kg) Planting sites should be cleared of competition to (Pfister 1974). improve shrub seedling success. Golden currant seed- Heit (1971) recommended that tetrazolium chloride lings and young plants are able to compete and estab- staining be used for testing viability, but embryo lish with some competitive understory. The shrub is excision is difficult. Seed viability is generally quite more sensitive to competition when planted along high, usually in excess of 75 percent. Seeds retain good streams or in moist situations, yet seedlings of this viability for 5 to 17 years if stored dry in sealed species are able to compete well with aggressive herbs containers (Pfister 1974) or in open warehouses that exist in riparian communities. New seedlings (Plummer and others 1968; Stevens and others 1981a). appear from natural seeding throughout native ranges, Seed may remain dormant in the soil for many years which attests to the survival attributes of this species. (Moss and Wellner 1953; Quick 1954).

690 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Chapter 23 Shrubs of Other Families

Bareroot or container-grown transplants can be recommended use of this species to reclaim coal fields quickly grown, although overseeding is usually recom- in south-central Utah. The species prefers fertile, mended to assure emergence of the desired number of well-drained, neutral, loamy soils (Dittberner and seedlings. Planting beds are normally thinned to the Olsen 1983; Wasser 1982). Haeussler and Coates proper spacing and density. One-year-old transplants (1986) reported the plant grows best on soils supplied develop a dense, well-branched root system, and field with humus. survival of 1-0 transplants is usually very high. Trans- Golden currant is widely used for hedges, wind- plants establish very well on harsh disturbances due, breaks, and conservation and landscape plantings in part, to the well-developed root system of young (Cook 1981; Wasser 1982). It is usually planted in plants. Container stock is grown from seed or hard- combination with other woody species, particularly wood or softwood cuttings (Doran 1957). evergreens or less open deciduous species. It is often planted as a conservation or wildlife habitat species, Uses and Management—Golden currant has been but other shrubs provide more acceptable forage for widely planted for wildlife habitat (Patton and Ertl birds (Miller and others 1948). This species is com- 1982), ground cover, watershed protection, and con- monly planted in rows or hedges to provide berries for servation plantings (Pfister 1974). It establishes with jam and jelly. good success in most wildland plantings; consequently, Golden currant is widely distributed along stream- it is often used to assure the establishment of a desir- banks, floodplains, and drainageways (Hitchcock and able species. others 1961; Wasser 1982). It can be used to reclaim Golden currant is a highly preferred spring and riparian disturbances, particularly moist sites where midsummer browse for big game, and is recommended the water table has been lowered by erosion and for planting in mountain brush, pinyon-juniper, big downcutting of gullies. sagebrush, and wet meadow communities that receive Golden currant is moderately fire tolerant (Hopkins more than 20 inches (51 cm) of annual precipitation and Kovalchik 1983; Rowe 1983), and recovers by (Plummer and others 1968; Wasser 1982). Kufeld and resprouting or natural seeding (Rowe 1983). Pfister others (1973) reported moderate summer use, but only (1974) reported that golden currant seedlings estab- light fall use by big game animals. The plant is eagerly lish well on mineral soils where the organic duff is grazed in early spring and summer, and can be planted removed by fire. Currants are subject to defoliation by on riparian sites and upland ranges to provide sea- the western tent caterpillar (Furniss and Barr 1975); sonal grazing by birds and big game. they are also an alternative host for white pine blister Golden currant is commonly planted with a combi- rust (Quick 1954). nation of other species to furnish diversity, early developing forage, fruits, and habitat for big game and Varieties and Ecotypes—None. birds. It provides preferred roosting and nesting cover for several songbirds (Johnson and Anderson 1980). Chukar partridge feed on the ripe fruits (Gullion Family Saxifragaceae ______1964). Dayton (1931) reported that this shrub species Ribes cereum has about average palatability for domestic livestock. Wax currant Although Dittberner and Olsen (1983) reported considerable variation in palatability among golden cur- Description—Wax currant (fig. 57) is a spreading, rant populations in the West, this species produces an unarmed, multicrowned, deciduous shrub, usually 2 to 5 excellent amount of herbage annually. ft (0.6 to 1.5 m) tall with puberulent and more or less Golden currant is an excellent species for stabiliza- stipitate glandular young branches (Hitchcock and oth- tion of roadways (Wasser 1982) and other distur- ers 1961; Welsh and others 1987). Leaves are fragrant, bances, particularly if transplant stock is used. Both clustered on short spurlike branches (Sampson and bareroot and container-grown transplants establish Jespersen 1963), usually shallowly three- to five-lobed. and survive well on most disturbances. Transplants The upper surface of the leaves is subglabrous to grow rapidly and furnish excellent ground cover in 2 to shiny, and the lower surface is waxy glandular dotted 3 years. If planted at close spacings, 3 to 4 ft (0.9 to (Stubbendieck and others 1986). The inflorescence is 1.2 m) apart, an effective cover can develop quickly. pubescent and sticky glandular. Two or three flowers Plants are reasonably adapted to disturbances where occur in short drooping clusters. Flowers are greenish portions of the topsoil remain or where the topsoil has white to pink (Welsh and others 1987). Fruits are dull to been mixed with the planting media. Disturbed areas bright red, glandular to glabrous, and usually between that lack good fertility and consist of coarse fragments 0.2 to 0.3 inch (6 to 8 mm) long. are less likely to sustain this shrub. However, golden Ecological Relationships and Distribution— currant is quite drought tolerant and can persist on Wax currant is found from British Columbia to south- harsh mine sites. Ferguson and Frischknecht (1985) ern California on the east slopes of the Cascade and

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Fruits are harvested by hand picking or flailing the bush to dislodge the berries. Fruits are macerated to remove the seed; the material is then dried, and seeds are separated from the dry pulp using a fanning mill. Seeds are quite small (Vories 1981), averaging about 251,000 per lb (553,400 per kg) (Pfister 1974). Seed production from wax currant is generally lower than from golden currant and sticky currant. In addition, seed germination is usually low and a longer period of wet prechilling is required to overcome dormancy. Pfister (1974) recommended that seeds be wet prechilled for 120 to 150 days at 28 to 32 ∞F (–2.2 to 0.0 ∞C). Seeds should be fall sown at a depth of 0.2 to 0.5 inch (6 to 13 mm) on a firm seedbed. Haeussler and Coates (1986) recommended seeding on moist min- Figure 57—Wax currant growing on a rocky out- eral soils with high percentages of organic matter. crop above timberline. Plummer and others (1968) reported that seeds of this species have low seed germination, and the initial establishment of new seedlings is only fair. Young plants grow slower than golden currant seedlings, but persistence of established plants is excellent. Plants Sierra Mountains, and eastward to Montana, Ne- spread well from natural seeding. It is less success- braska, and New Mexico (Hitchcock and others 1961). fully established by transplanting than golden cur- It is one of the most widely distributed of all western rant. Wax currant is easily cultured in nursery beds. currants, because it occurs from sagebrush deserts to Seeds of wax currant may be spread by birds or small alpine areas (Great Plains Flora Association 1986; mammals (Kramer 1984) or by being deposited di- USDA Forest Service 1937). Ribes cereum var. cereum rectly below the shrub (Moss and Wellner 1953). Seeds is found from British Columbia to southern Arizona may remain viable for a long period (Lyon and Stickney and southern California, and east to central Montana, 1976). Pfister (1974) reported seeds retained good Idaho, and western Nevada. The R. c. var. inebrians is viability after 27 years of storage. more common eastward from central Idaho and Mon- Although seeds are relatively thin walled and can tana, south and east to Nebraska, New Mexico, Utah, be destroyed by severe fires (Kramer 1984), they and eastern Nevada. Welsh and others (1987) re- benefit from fire scarification (Morgan and ported that most plants in Utah are R. c.var. inebrians, Neuenschwander 1985; Young 1983). New seedlings but intergradation of morphological features is com- often appear following low-severity fires that remove mon, and no geographical correlation is apparent in little surface litter (Rowe 1983). Plants have large Utah. R. c. var. colubrinum is confined to the Snake and deeply buried root crows that resprout from River Canyon and related tributaries in Idaho, Wash- belowground tissue if not destroyed by fire (Bock and ington, and Oregon (Hitchcock and others 1961). Bock 1984). Goodrich and Neese (1986) reported the plant is common and widespread in mountainous communi- Uses and Management—Wax currant is some- ties between 6,500 and 11,000 ft (2,000 to 3,400 m) in what less palatable throughout the entire year than Utah. Sampson and Jespersen (1963) reported that golden currant (Plummer and others 1968). Kufeld wax currant is more widely distributed in California and others (1973) reported that big game made only than sticky currant, although their distributions over- light use of the shrub in the spring and fall, but lap. Wax currant is common in various plant commu- Plummer and others (1968) reported it was heavily nities, but not as a dominant species. It is often browsed during all seasons. associated with antelope bitterbrush in the Central Palatability of wax currant is low for livestock, but Rocky Mountains (USDA Forest Service 1937). It is it is an important browse due to its abundance, pro- prevalent in various ponderosa pine/Douglas-fir forest ductivity, and availability (Dittberner and Olsen 1983; types in Colorado and several lodgepole pine associa- Mueggler and Stewart 1980; Sampson and Jesperson tions in Oregon (Volland 1985b). It occurs in aspen 1963). This shrub has been principally used for wild- communities (Mueggler 1988) and conifer forests of life habitat improvement, although limited plantings Utah (Youngblood and Mauk 1985) and Idaho (Steele have been established for watershed and ground- and Geier-Hayes 1987), but only as a minor species. cover protection. It is an important shrub for use in Plant Culture—Wax currant plants flower from restoration of mountain brush, aspen, and subalpine April to June; fruits ripen in August (Pfister 1974). communities (Plummer and others 1968).

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Wax currant has not been widely used for control of 12 flowers occuring in a cluster. Berries are 0.4 to severe erosion problems or disturbed land plantings. 0.5 inch (10 to 13 mm) long, black, rather dry, and It has been used to stabilize watersheds, small gullies, glandular (Welsh and others 1987). and moist streambanks in aspen and conifer forests if Ecological Relationships and Distribution— topsoil is present. Plants are adapted to disturbances This species is common from British Columbia to related to fire and logging. Natural regeneration California on the east side of the Cascade and Sierra frequently occurs after these disturbances, and this Nevada Mountains, and extending east to Montana species provides considerable ground cover and herb- and Arizona (Hitchcock and others 1961; Sampson age. It can be seeded and transplanted on areas ex- and Jespersen 1963; Welsh and others 1987). Hitchcock posed by logging or fires with acceptable success. The and others (1961) divided it into two intergrading shrub grows well with understory herbs, and can be geographic races. R. v. var. hallii is distinguished by planted with a number of species to control erosion and its glabrous ovaries and glaucous berries; it occurs protect unstable sites. It does not spread rapidly to throughout the eastern Cascades south of Mt. Rainier occupy harsh disturbances, and has not performed well and the Sierras. R. v. var. viscosissimum has glandu- as a pioneer species on mines, roadways, or similar lar and more or less pubescent ovaries and nonglaucous disturbances. This shrub grows well with some over- berries; it occurs over the remainder of the species’ story trees, and can be used to plant campgrounds and range. Welsh and others (1987) concluded that plants recreation sites where a dense tree cover may be de- occurring in Utah are unique; the herbage has dense sired. This species could be more widely used to restore stipitate-glandular and nonglandular hairs and long, native communities. broad hypanthia. Varieties And Ecotypes—None. Sticky currant grows in shady woods and rocky places in the Sierra Nevada Mountains at elevations between 6,000 and 9,000 ft (1,800 and 2,700 m) Family Saxifragaceae ______(Sampson and Jespersen 1963). Goodrich and Neese Ribes viscosissimum (1986) reported the species is common throughout the Sticky currant Uinta Basin in Utah, existing with aspen and conifer woods at 7,500 to 10,000 ft (2,300 to 3,000 m). Welsh Description—Sticky currant (fig. 58) is a spread- and others (1987) reported that this shrub occurs ing, unarmed, uneven, aromatic shrub, mostly 3.3 to throughout most of north, central, and eastern Utah, 6.6 ft (1.0 to 2.0 m) tall. Stems, leaves, inflorescences, growing in the shade of aspen, fir, Douglas-fir, lodge- and fruits are sparsely to thickly glandular (Welsh pole pine, and spruce woodlands, but is less common in and others 1987). Leaf blades are orbicular, three- to mountain brush and open meadows. Steele and Geier- seven-lobed with the main lobes crenate or dentate, Hayes (1987) reported sticky currant as a principal round in outline, heart shaped at base, and hairy or species with grand fir in central Idaho where it is often glandular on both surfaces (Sampson and Jespersen an early seral shrub, and may be the first shrub to 1963). Flowers are green, white, or pink, with three to dominate scarified sites following logging. These authors concluded that plants have a low tolerance for shade and begin to decline as a canopy taller than their own develops. Sticky currant plants attain mature stature within 5 years following logging. Their canopy is sparse, and the shrub does not seriously compete with the establishment of tree seedlings (Steele and Geier-Hayes 1987). However, plants of sticky currant remain longer than some species of ceanothus as overstory shade develops. Sticky currant is not a principal species in any conifer forest (Youngblood and Mauk 1985) or aspen type (Mueggler 1988) in Utah. Plant Culture—Sticky currants flower in May or June, and fruits ripen in August or September (Pfister 1974). Fruits are harvested by hand picking or beating the bush to dislodge the berries onto canvas or other collection tarps. Fruits are cleaned by maceration using a Dybvig separator. Filled seeds will sink or settle in water, and the pulp and empty seeds can be Figure 58—Sticky current generally occurs above decanted (Pfister 1974). Following maceration, large timberline on windswept ridges.

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 693 Chapter 23 Shrubs of Other Families seedlots can also be dried and the seeds separated topsoil is disrupted or removed. The species is more from the debris using fanning mills (Plummer and site specific than other shrubs in its genus. Planting others 1968). There are 255,000 to 349,000 cleaned unadapted ecotypes on disturbances reduces survival; seeds in 1 lb of cleaned seed (562,000 to 769,000 per kg) however, site factors affecting adaptation are not well (Pfister 1974). understood. Most plants produce some fruits each year. New or The plant can be used as a nurse crop to improve young stands produce the most consistent crops. seedling establishment of tree seedlings. It is also Although fruits are large, seed collection is slow and relatively abundant in localized areas and should be costly. Seed costs and poor availability discourage the maintained for game habitat and ground cover. This use of this species in large projects. Enough seeds can species frequently increases following logging and be economically harvested most years from specific becomes important for watershed protection. locations to supply seeds for rearing transplant stock. Varieties and Ecotypes—None. Seeds can be stored for extended periods, exceeding 17 years (Pfister 1974), if kept in dry sealed containers. Steele and Geier-Hayes (1987) reported seed buried in Family Scrophulariaceae ______soil and duff remain viable long after the parent shrubs disappear. Penstemon fruticosus Seeds require a wet prechilling period of 140 days at Bush penstemon 28 to 32 ∞F (–2.2 to 0 ∞C) to initiate germination (Pfister 1974). Seed germination is only moderate. Seeds gath- Description—Bush penstemon is one of approxi- ered from different collection sites can exhibit variable mately 230 species of Penstemon that occur in Western germination patterns. Plummer and others (1968) North America (Hylton 1974). Most are herbaceous reported that initial establishment was only fair plants, but the list includes a few suffrutescent or because of poor germination and relatively slow growth woody shrubs. Bush penstemon is a woody species that of small seedlings. Growth of young plants is some- normally reaches a height of 2.5 ft (76 cm). Plants form what slow, but improves as plants attain mature a persistent, woody crown that is usually wider than stature. However, plants that reach 2 to 3 years of age tall. They develop into wide, dense clumps when young may succumb under adverse conditions; final estab- and spread by stem layering, but natural dieback lishment of seeded plots is only moderate or good. leaves the center open. Stems are erect, and clusters of Plants are not rhizomatous, and natural spread is leaves form at the base of current year’s stems (Hitchock slow unless sites are cleared by logging, fires, or and Cronquist 1976). Leaves are mostly lustrous, related disturbances. evergreen, lanceolate or oblanceolate to elliptic, en- This species can be produced as bareroot nursery tire or serrulate or denticulate, 0.4 to 2 inches (1 to stock or container material. Initial establishment is 5 cm) long, and 0.2 to 0.6 inch (5 to 15 mm) wide. poor even under controlled rearing conditions; re- Flowers are bright lavender blue, 1 to 1.5 inches (25 planting is often required to produce full stands. Once to 38 mm) long, 0.3 to 0.5 inch (7 to 12 mm) wide (Davis established, nursery stock grows well, and when trans- 1952). Three varieties were described by Hitchcock planted to range or wildland sites, survives quite well. and Cronquist (1973); they differ in regard to leaf length, width, margins, and size of flowers. Penstemon Uses and Management—Sticky currant is recom- fruticosus var. serratus is mainly confined to the Snake mended for planting aspen, Douglas-fir, spruce woods, River Canyon of Oregon and Idaho; P. f. var. scouleri and less commonly mountain brush communities occurs in northeastern Washington, northern Idaho, (Plummer and others 1968). It has been used to en- and adjacent British Columbia; and P. f. var. fruticosus hance wildlife habitat (Pfister 1974; Plummer and is more widespread and variable, but generally does others 1968). Plants have only fair or poor palatability, not occupy the range of the two other varieties but are quite tolerant of grazing. In Utah the plant is (Hitchcock and Cronquist 1973). most commonly used during spring and summer peri- Ecological Relationships and Distribution— ods by big game (Plummer and others 1968); heavy Bush penstemon generally occurs at elevations over browsing may occur during these periods. Sampson 4,000 ft (1,200 m) on rocky slopes in forest openings and Jespersen (1963) reported that plants in Califor- from the Cascades of Washington and Oregon across nia produce abundant herbage, which is heavily Idaho to the Rocky Mountains of Montana and cropped by sheep and deer, particularly in fall. Wyoming (Davis 1952). Although widespread, this Sticky currant grows well with other species, and species is not particularly abundant. It is not a princi- frequently occurs with many other shrubs and herbs. pal understory with any forest community in the It can be planted to stabilize disturbances following Intermountain region (Mueggler 1988), northern Idaho fire and logging. It grows well on undisturbed soils, but (Cooper and others 1987), or eastern Washington is less vigorous and poorly adapted on sites where the

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(Daubenmire and Daubenmire 1968). Plants normally opens and seeds shatter. Sites that are good seed occur on ridge crests, talus slopes, and forest openings. producers have remained so for nearly 20 years. Most They invade roadcuts and are noticeable on aban- plants within a stand produce good seed crops each doned disturbances. Bush penstemon is encountered year. Seeds are easily harvested and can be cleaned by in scattered stands of ponderosa pine and Douglas-fir. allowing the capsules to dry and dehisce, releasing the At higher elevations it exists in openings with subal- seeds. Seeds are small, but can be separated from the pine fir. It does not occur in dense shrub fields or mixed debris using a fanning mill or gravity table. grass/herb communities. Some seeds will germinate soon after harvesting if exposed to favorable conditions, but a period of Plant Culture—Bush penstemon usually occurs as afterripening and wet prechilling is normally required scattered plants on rocky outcrops (fig. 59). Following to assure uniform germination. Seeds that are fall disturbances or clearing of overstory species, this seeded receive natural wet prechilling and germinate shrub flourishes. It invades openings and forms patches uniformly in spring. Bush penstemon seedlings are dominating small rocky areas. Young plants are vigor- vigorous and normally grow rapidly. ous and robust. Disturbances often become excellent New seedlings spread quickly from existing plants. seed production centers. Although seedlings increase rapidly, the young plants Plants are evergreen; a high percentage of leaves are not highly competitive with grasses and broadleaf remain on the shrub throughout the winter. New herbs. New seedlings are well adapted to harsh sites growth begins early in spring before most other plants and compete well with other species under these initiate growth. Leaves develop rapidly to form a dark conditions, but they are not able to compete with green clump. grasses and broadleaf herbs on deep, fertile soils. Plants usually produce a profusion of flowers each Seedlings establish well by broadcast seeding on a year. Flowering begins in early June and may continue rough seedbed or by drill seeding at shallow depths, for nearly 1 month. Seeds ripen from mid-August to about 0.24 inch (6 mm) on a firm seedbed. Seed late September. Numerous seeds develop each year, germination and establishment occur uniformly; er- but not all are viable. Seed quality varies widely ratic stands are uncommon. among collection sites and years. Although the plant Bush penstemon is well adapted to drill or broadcast flowers for an extended period, seeds ripen fairly seeding. The seeds can be planted separately or in uniformly and remain in the capsule for nearly a mixtures with most drills or broadcast seeders. The month before shattering. seeds are small and need not be planted at rates Seeds are collected by hand stripping the dry cap- exceeding 2 to 3 lb per acre (2.2 to 3.4 kg per ha). sules, or by clipping the inflorescences. They must be Seeding at a rate of 2 to 4 lb per acre (2.2 to 3.4 kg per harvested soon after the capsule dries, but before it ha) is recommended for drill seeding. Broadcast seeding on unprepared road disturbances may be increased to 4 to 6 lb per acre (4.5 to 6.7 kg per ha), depending on whether other species are seeded and the condition of the seedbed. This is one species that responds well to the amount of seed sown. Increasing the seeding rate usually results in an increase in the number of plants to establish. Thus, plant density of this species can be regulated by adjusting the seeding rate. If seeded concurrently with species having larger seeds, it is recommended that the smaller seeds of bush penstemon be planted in separate drill rows to better regulate planting depth. Small seedlings are quite vigorous. When seeded in barren openings, bush penstemon matures within 2 to 3 years. When grown in competition with grasses, the young plants can be seriously stunted. However, bush penstemon can be planted with certain grasses and forbs. Germination and seedling emergence of this shrub occur at about the same time as most herbs. Consequently, bush penstemon seedlings are able to compete favorably with many native grasses and broad- Figure 59—Bush penstemon, a low-growing mound- leaf forbs. Bush penstemon seedlings are not able to ing shrub, generally grows on rocky outcrops. persist when seeded with a dense cover of aggressive

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 695 Chapter 23 Shrubs of Other Families grasses, including smooth brome, intermediate wheat- considerable usefulness for formal plantings in land- grass, or timothy. scaping commercial or residential sites. Its evergreen Bush penstemon grows quickly from direct seeding growth habit suggests this native shrub would be or transplanting. Transplants can be reared as bareroot particularly useful for planting around summer homes nursery stock, greenhouse-grown container stock, or and dwellings to prevent the spread of wildfires. rooted stem cuttings. This species is also an important plant for wildlife Bush penstemon seedlings begin growth early in habitat. It occupies sites that are open and exposed spring, often before nursery lifting is completed. If during midwinter. Game animals are often confined to bareroot stock initiates growth prior to lifting, field these sites and rely heavily on this shrub. It has survival of the transplants declines significantly. Fall considerable promise for many situations, and could lifting and overwinter storage may be most practical. be more widely used in restoration programs if seeds Plants stored in bags or crates should have the tops were more available. exposed. Plants can be fall lifted and planted in saw- Varieties and Ecotypes—None. dust or other media for overwinter storage in open lath houses. Bareroot stock transplants very well and can provide effective ground cover within 1 to 2 years. Family Ulmaceae ______Bush penstemon is easily grown from seed or stem cuttings. However, plants grow rapidly and can be Celtis reticulata easily stunted if reared in containers that are too Netleaf hackberry small. Container stock grows best in a well-drained, neutral potting media. Stem cuttings taken in mid- Description—Celtis is a large genus of about 70 winter and early spring root readily. species of shrubs and trees of the Northern Hemi- sphere. Netleaf hackberry is a variable taxon, and Uses and Management—Bush penstemon is well includes varieties that are difficult to separate due to adapted for revegetation of road disturbances, mine the many intergradations encountered (Harrington sites, and other drastic disturbances (Hungerford 1964). Netleaf hackberry, also known as hackberry, 1984; Monsen and Christensen 1975; Plummer 1977; western hackberry, and palo blanco, is a large decidu- Tiedemann and others 1976). From roadway plantings ous shrub to small globose tree growing to 30 ft (9.1 m) in northwestern Montana, Hungerford (1984) rated in height and 6 to 12 inches (15 to 30 cm) in diameter, bush penstemon as the second most useful species for depending on available water (Bonner 1974a; Hitchcock revegetation of forest roads. Only Woods’ rose ex- and others 1973). Crowns are open and irregular with ceeded this species in overall usefulness. The plant is stout, ascending branches. Young twigs are glabrous extremely vigorous, matures quickly, and spreads to pubescent. Bark of older branches is reddish gray rapidly by natural seeding and crown spread with prominent ridges and furrows. Plants develop a (Hungerford 1984; Monsen and Christensen 1975). It spreading and fibrous root system. Leaves are short provides excellent ground cover due to its dense, low- petiolate and leathery with scabrous surfaces and profile growth habit. It furnishes effective protection entire to serrate margins. They are ovate or ovate- throughout the entire year, as plants are evergreen, lanceolate, usually somewhat acuminate and acute to and provides aboveground protection at all seasons. It chordate at the base, 1 to 4 inches (2.5 to 10.2 cm) long is able to persist and spread when buried by shifting and 0.74 to 1.5 inches (1.9 to 3.8 cm) wide. Leaf soil. It is one of few species that is well adapted to both surfaces are dull green to yellow green, but paler with rocky, shallow soils, as well as deep, well-developed conspicuous reticulate veinlets below. They are three soils. This species can be successfully seeded or trans- nerved near the base and short pilose along the veins. planted on rocky roadcuts. It establishes very well Foliage turns yellow in fall (Hitchcock and others from broadcast seedings with minimal soil coverage. 1973; Welsh and others 1987). Few other species, including most grasses or broadleaf Plants are polygamo-monoecious with greenish flow- herbs, establish as successfully from broadcast seeders that emerge with the leaves clustered on short ing as this shrub. shoots of the current season. Flowers develop on slen- Bush penstemon is an attractive shrub that flowers der, pubescent pedicels. Lower flowers are mostly throughout most of the summer and provides an at- staminate and several per axil. Upper flowers are one tractive leafy cover. It is able to spread and occupy or two per axil and mostly perfect, but some may be rocky sites, providing an interesting pattern of ground pistillate. Perfect flowers consists of five membranous cover. Both the flowers and leaves furnish an attrac- perianth segments, five stamens, and a one-seeded tive background for other species. The shrub can be ovary with spreading style tips. Fruit is an orange-red used to treat roadways, mine sites, and recreational to yellow, thin-walled, globose drupe with a datelike areas where aesthetic values are important. It has flavor (Treshow and others 1970).

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Ecological Relationships and Distribution— water apparently aids in improving germination. Netleaf hackberry grows on dry foothills, open slopes, Bonner (1974a) reported that fermenting the fruits for rocky bluffs, watercourses, and canyons from the Snake 3 days at room temperature prior to depulping and wet River drainage of the Northwestern States, east to prechilling improved germination. Approximately Colorado, and south to west Texas and southern Cali- 80 lb (36.3 kg) of seed may be recovered from 100 lb fornia (Bonner 1974a; Preston 1968; Thornburg 1982). (45.4 kg) of fruits; there are about 4,870 cleaned seeds It grows at elevations ranging from 2,000 to 6,000 ft per lb (10,736 per kg) (Swingle 1939). Dry fruits or (610 to 1,800 m) (Sutton and Johnson 1974; Thornburg cleaned seed may be stored in sealed containers at 1982) on well-drained soils ranging from silty to rocky, 41 ∞F (5.0 ∞C). Seeds require either fall sowing or a shallow to deep, and neutral to calcareous (Sutton and 120-day laboratory wet prechilling at 41 ∞F (5.0 ∞C) Johnson 1974; Thornburg 1982; Van Dersal 1938). prior to spring sowing in the nursery (Hartmann and Plants grow in full sunlight and are drought tolerant others 1990). and wind resistant (fig. 60). Netleaf hackberry is well Germination is usually quite low as seed from most adapted to watercourses and canyons where it may wildland collections has poor fill (Bonner 1974a). Un- grow on limestone and gravelly and rocky soils developed seeds are difficult to separate and remove (Thornburg 1982). during cleaning. Netleaf hackberry occurs as scattered plants in big Seedlings grow slowly and are sensitive to competi- sagebrush and mountain brush communities at 3,000 tion from herbs. Nursery grown transplants and rooted to 5,000 ft (910 to 1,500 m) in Utah (Welsh and others cuttings are often recommended for field plantings to 1987). It is prevalent on steep south and west slopes reduce problems associated with establishment from with pinyon-juniper, antelope bitterbrush, Stansbury seed. Nursery or container-grown stock develops well, cliffrose, and other mountain brush species through- and field survival is excellent. One or 2-year-old plant- out the Wasatch Mountains. It is particularly impor- ing stock is recommended for outplantings. tant on exposed slopes with other persistent species. Seed and planting stock are rarely available on the commercial market; thus, it is necessary to arrange for Plant Culture—Flowering occurs in March and collection of seed or propagation of seedlings. Al- April; fruits ripen in late fall, and seed disperses in fall though extensive testing of different ecotypes has not and winter (Swingle 1939). Fruits persist on the trees been reported, considerable differences in vegetative for several months after ripening, providing a pro- traits and areas of occurrence would suggest that longed period for collection. Although this plant often planting success is dependent on the use of an adapted grows in harsh environments, some seeds are usually seed source. produced every year. They are gathered by hand or by knocking them onto tarps. Collection may be easier in Uses and Management—Netleaf hackberry cur- winter after leaves have fallen (Bonner 1974a). Drying rently receives limited use in artificial plantings. How- may not be required if seeds are collected late in the ever, it is one of the tallest native species on sites season, but seeds are more difficult to remove from dry within its range of adaptation, and provides important fruits. Twigs and other debris are removed by fanning. habitat for wildlife. It occupies exposed windblown Extraction of the pulp by macerating the fruits in slopes where big game animals concentrate, providing important cover and protection. The shrub is considered fair to good forage for deer and livestock (Sampson and Jespersen 1963). It frequently grows as clumps or in groups to furnish pockets of cover where few other shrubs are able to grow. The species grows as a border along riparian zones and significantly enhances these communities. It is particularly adapted to steep slopes and well-drained sites aligning stream bottoms. Fruits persist on the tree well into the winter and provide food for birds (Thornburg 1982). Montezuma quail reportedly eat the seeds during winter months (Bishop and Hungerford 1965). Gambel quail also forage on the fruits (Gullion 1964; Martin and others 1951), as do various small mammals (Bailey 1936). This species has established well from plantings on big game ranges in central and south-central Utah Figure 60—Netleaf hackberry growing on a dry, open when planted on shallow soils and exposed slopes slope protected by a rocky outcrop. within the lower mountain brush zone. It has been

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 697 Chapter 23 Shrubs of Other Families planted in groups or patches to provide concealment summer storms may occur. Plants are generally very and winter cover in areas where upright vegetation is hardy and long lived. Twigs and leaves are frequently required. Game animals and livestock may graze new infested by insect galls (Welsh and others 1987), but plantings, but not seriously enough to adversely affect vigor is not seriously affected. Plants are not especially plant growth. This species has survived years of drought fire tolerant, and can be killed by wildfires, although without appreciable reduction in plant size and vigor. resprouting does occur. Many native stands persist on Once established, the young plants are very hardy and rocky sites that provide some protection from wild- persistent. They are able to compete and grow with fires. Plants that do survive fires usually recover understory herbs. slowly. Plants do not spread rapidly by natural seed- Netleaf hackberry can be used to provide shade in ing. Few new seedlings appear following fires or other recreation sites, and for windbreaks, shelterbelts, or disturbances. Seedlings appear to require shading, domestic landscaping. It can be used on sites receiving but seedbed requirements are not well understood. at least 14 to 16 inches (36 to 41 cm) of annual precipi- Improved Cultivars and Other Species—None. tation. It does not require maintenance in landscape In the Midwest, “Oahe,” a released cultivar of Celtis plantings, and could be more widely used in these occidentalis, has been used as substitute for Siberian situations. elm, Russian-olive, and green ash, although vigor and Young plants do not grow rapidly. More rapidly ease of establishment are lower (Soloman 1985). The developing species are recommended for unstable cultivar has received little testing in the Intermoun- watershed or mine disturbances. This plant provides tain region. important ground cover on sites where high-intensity

698 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Spreader 451 Spreader II 451 Index Teton 451 Travois 451 alfalfa, range type 427 A alfalfa, sicklepod 447Ð448 alfileria 291, 427 Acer glabrum See maple, Rocky Mountain Allenrolfea occidentalis See iodine bush Achillea millefolium lanulosa See yarrow, western Alnus incana See alder, thinleaf Achnatherum hymenoides See ricegrass, Indian Alopecurus arundinaceus See foxtail, creeping Achnatherum thurberianum See needlegrass, Thurber Alopecurus pratensis See foxtail, meadow acid equivalent 97 Alopecurus ventricosus See foxtail, creeping aerial broadcasting 77, 205 Amelanchier alnifolia See serviceberry, Saskatoon aerial ignitors 85 Amelanchier utahensis See serviceberry, Utah aerial seeding 77Ð79, 205 ammonium 54 fixed-wing aircraft 78 anchor chain 68Ð69 helicopter 78Ð79 disk-chain 69 aesthetics 133 Dixie sager 69Ð70 Agropyron cristatum See wheatgrass, crested; wheat- Ely 66, 68, 70 grass, crested fairway smooth 66 Agropyron cristatum x A. desertorum See wheatgrass, swivels 231Ð232 crested hybrid Andropogon curtipendula See grama, sideoats Agropyron dasystachyum See wheatgrass, thickspike angelica, small leaf 269, 427 Agropyron desertorum See wheatgrass, crested animal control See depredation standard animal depredation See depredation Agropyron elongatum See wheatgrass, tall annuals, spring-growing 257 Agropyron fragile See wheatgrass, crested Siberian annuals, summer-growing 258 Agropyron inerme See wheatgrass, bluebunch; Apache plume 253, 273, 276, 293, 559, 561Ð563, 601, wheatgrass, bluebunch bear 704, 724, 748 Agropyron intermedium See wheatgrass, intermediate Arctostaphylos uva-ursi See manzanita, bearberry Agropyron repens x A. spicatum See wheatgrass, Aristida purpurea See threeawn, red hybrid: NewHy Aroga websteri See moth, sagebrush defoliator Agropyron sibiricum See wheatgrass, crested Siberian Arrhenatherum elatius See oatgrass, tall Agropyron smithii See wheatgrass, western Artemisia abrotanum See wormwood, oldman Agropyron spicatum See wheatgrass, bluebunch Artemisia arbuscula See sagebrush, low Agropyron spicatum x A. dasystachyum See wheat- Artemisia bigelovii See sagebrush Bigelow grass, hybrid: SL1 Artemisia cana See sagebrush, silver Agropyron trachycaulum See wheatgrass, slender Artemisia filifolia See sandsage Agropyron trichophorum See wheatgrass, intermediate Artemisia frigida See sage, fringed Agrostis alba See bentgrass, redtop Artemisia longiloba See sagebrush, alkali Agrostis palustris See bentgrass, redtop Artemisia ludoviciana See sage, Louisiana Agrostis stolonifera See bentgrass, redtop Artemisia nova See sagebrush, black air-screen separator 715 Artemisia pygmaea See sagebrush, pygmy Aira cespitosa See hairgrass, tufted Artemisia rigida See sagebrush, stiff alder, thinleaf 266Ð267, 270, 272, 601, 616Ð618, 748 Artemisia spinescens See budsage alfalfa 132, 135, 136, 138, 142, 182, 183, 196, 206, Artemisia tridentata See sagebrush, big 216, 220, 237, 242, 259, 264, 269, 271, 273, 275, Artemisia tripartita See sagebrush, threetip 277, 280, 282, 284, 287, 289, 425, 448, 449Ð450, artificial seeding 516, 539 702, 727, 741 ash, singleleaf 276, 278, 601, 650Ð652, 704, 748 Drylander 451 aspen 119, 195, 214, 266Ð267, 748 Ladak 450Ð451, 724, 728 aspen communities 118, 215, 308Ð309 Nomad 450, 724 aspen low forb 118 Rambler 450 aspen shrub 118 Rhizoma 451 aspen tall forb 118 Runner 451 aspen-conifer 195, 213, 269

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-1 mixed forb 118 barley, meadow 259, 261, 264, 265, 269, 289, 308, mixed shrub 118 311 aspen-conifer communities 9, 216, 601 bassia, fivehook 264 aspen openings 204Ð205 bentgrass, carpet 310 See also bentgrass, redtop aspen openings, upper elevation 201, 259 bentgrass, creeping 310 See also bentgrass, redtop aster 196, 202, 223, 237, 246Ð248, 252, 255Ð256, bentgrass, redtop 208, 259, 261, 264, 265, 269, 308, 259, 261Ð262, 264, 269, 271Ð272, 275, 277Ð278, 310, 345Ð346, 700 280, 282, 289, 294, 741 Streaker 298, 346 aster, alpine leafybract 259, 261 Betula glandulosa See birch, bog aster, blueleaf 135, 261, 269, 271, 277, 289, 426Ð427, Betula occidentalis See birch, water 433, 702, 724 Betula papyrifera See birch, paper Aster chilensis See aster, Pacific big game depredation See depredation aster, Engelmann 269, 289, 427, 702, 724 big sagebrush-cheatgrass 106 Aster glaucodes See aster, blueleaf bighorn sheep 165Ð166 aster, Pacific 259, 261Ð262, 264, 269, 271, 275, 277, biological insect control 721 280, 282, 294, 426Ð427, 431Ð432, 702, 724, 763 birch, bog 601, 618Ð620 aster, smooth 259 birch, paper 601, 622, 748 Astragalus cicer See milkvetch, cicer birch, water 262, 266, 267, 601, 620Ð621 astragalus, giant 731 bitterbrush, antelope 103, 132, 135, 138, 142, 143, Atriplex buxifolia 481Ð482 See also saltbush, Gardner 181, 182, 183, 185, 196, 270, 272, 273, 276, 278, Atriplex canescens See saltbush, fourwing 280, 283, 288, 557, 559, 581Ð586, 587, 601, 704, Atriplex confertifolia See shadscale 724, 727, 730, 741, 748 Atriplex corrugata See saltbush, mat Lassen 586 Atriplex cuneata See saltbush, Castle Valley clover bitterbrush, desert 273, 276, 293, 559, 579Ð581, 582, Atriplex falcata 481Ð482 See also saltbush, Gardner 601, 704, 724, 727, 748 Atriplex gardneri See saltbush, Gardner blackbrush 195, 199, 232, 246, 247, 252, 253, 254, Atriplex hymenelytra See saltbush; saltbush, desert 257, 258, 293, 294, 552Ð555, 601, 704, 748 holly blackbrush communities 9, 252Ð253, 293, 308, 601 Atriplex lentiformis See saltbush; saltbush, quailbush blackcap 751 See raspberry, black Atriplex obovata See saltbush; saltbush, broadscale blackthorn 577 Atriplex polycarpa See saltbush; saltbush, allscale bladersenna, common 741 Atriplex tridentata 481Ð482 See also saltbush, bluebell 741 Gardner bluebell, tall 259, 269, 427, 452, 453 auger 765 bluegrass 102 Avena elatior See oatgrass, tall bluegrass, alpine Gruening 298 B bluegrass, big 228, 259, 269, 271, 275, 277, 282, 289, Balsamorhiza hookeri See balsamroot, hairy 308, 401Ð402, 700 Balsamorhiza macrophylla See balsamroot, cutleaf Sherman 298, 402 Balsamorhiza sagittata See balsamroot, arrowleaf bluegrass, bulbous 311 balsamroot, arrowleaf 132, 135, 138Ð139, 142, bluegrass, Canada 259, 269, 271, 275, 277, 282, 289, 182Ð183, 185, 196, 271, 275, 277, 282, 287, 308, 311, 402Ð404, 700 426Ð427, 435Ð436, 702, 724, 727Ð728, 741 Foothills 298, 404 balsamroot, cutleaf 271, 277, 287, 426Ð427, 436, 702, Reubens 298, 404 724, 727, 741 bluegrass, Cusick’s 311 balsamroot, hairy 271, 277, 426Ð427, 436 bluegrass, Kentucky 196, 269, 308, 311, 406Ð408, 724 barberry, creeping 601, 704, 748 See also Oregon Newport 138, 298, 408 grape bluegrass, longtongue 311 barberry, Fremont 614, 704 bluegrass, mutton 308, 311, 404Ð406 barberry, shining 613 bluegrass, nodding 311 bareroot stock 739Ð742, 747, 754Ð755, 763Ð767 bluegrass, pine 311 barley, beardless 311 bluegrass, Sandberg 135, 182Ð183, 185, 264Ð265, barley, bulbous 308, 311, 700 273, 275, 277, 279, 280, 282, 284Ð287, 291Ð292, barley, common 311 294, 308, 311, 408Ð410, 700 barley, foxtail 269, 311 Canbar 298, 410

Index-2 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 High Plains germplasm 298, 410 brushland plow 66 Service 298, 410 buckthorn, cascara 266Ð267, 601, 704 boron 51, 52, 54 buckwheat 652Ð653 See also eriogonum bouncing-bet 427 buckwheat, wild California flattop 704 Bouteloua curtipendula See grama, sideoats budsage 245Ð246, 291, 495, 511Ð513 See also Bouteloua eriopoda See grama, black sagebrush, budsage Bouteloua gracilis See grama, blue budsage communities 245 boxelder 602 buffaloberry, roundleaf 273, 293, 601, 643Ð644, 704, Brillion Seeder 80 748 broadcast ground application 93 buffaloberry, russet 601, 642Ð643, 704, 748 broadcast seeder 72, 76Ð77 buffaloberry, silver 262, 266Ð267, 601, 641Ð642, 704, broadcast seeding 76, 141 731, 748 broadcast spray application 92 bulrush, saltmarsh 263 brome, California 310 See also brome, mountain bulrush, tule 263 Cucamonga 298 Bureau of Plant Industry 2, 3 brome, cheatgrass See cheatgrass burnet, small 132, 135, 138, 142, 182Ð183, 185, 196, brome, downy See cheatgrass 271, 273, 275, 277, 279, 280, 282, 284, 286, 287, brome, erect 310 294, 425, 427, 457Ð459, 702, 724, 728, 741, 763 brome, foxtail 310 Delar 459 brome, fringed 358Ð360 burning, prescribed 101, 106, 160, 168 brome, Hungarian See brome, smooth aspen communities 118Ð119 brome, meadow 259, 264Ð265, 269, 289, 310, cheatgrass 116 365Ð367, 700 effects 105 Fleet 367 pinyon-juniper 116Ð117 Paddock 367 postfire management 119 Regar 196, 269, 271, 275, 277, 282, 287, 298, 310, sagebrush 115Ð116 367, 724, 728 uses 101 brome, mountain 135, 138, 196, 259, 261, 264, 265, 269, 271, 289, 308, 310, 360Ð362, 700, 724 C Bromar 298, 362 cables 67, 68, 69 See also anchor chain Garnet 362 Calamagrostis canadensis See reedgrass, bluejoint brome, nodding 308, 310, 358Ð360 calcium 51, 54, 178Ð179 brome, red 254, 255, 310 calcium nitrate 54 brome, red communities 9, 253Ð256, 294 camphorfume, Mediterranean 469, 491 brome, smooth 132, 135, 138, 142, 182Ð183, 185, Camphorosoma monspeliaca See camphorfume, 196, 205, 259, 261, 264Ð265, 269, 271, 275, 277, Mediterranean 282, 287, 289, 310, 363Ð366, 700, 727, 728, 731 canarygrass, reed 135, 182Ð184, 196, 261, 308, 311, Achenbach 299 396Ð398 Lincoln 299, 365 Ioreed 299, 397 Manchar 299, 365 cattle 176, 178Ð179 northern 259, 308 Ceanothus 656 Polar 299 Ceanothus cuneatus See ceanothus, wedgeleaf southern 308, 724 ceanothus, deerbrush 601, 658Ð659, 704, 748 varieties 365 ceanothus, desert 657Ð658 brome, subalpine 259, 269, 289, 308, 700 ceanothus, Fendler 273, 276, 601 Bromus anomalus See brome, nodding Ceanothus fendleri See ceanothus Fendler Bromus biebersteinii See brome, meadow Ceanothus greggii See ceanothus, desert Bromus carinatus See brome, California; brome, Ceanothus integerrimus See ceanothus, deerbrush mountain ceanothus, Lemmon 659 Bromus ciliatus See brome, fringed Ceanothus lemmonii See ceanothus, Lemmon Bromus erectus See brome, meadow ceanothus, Martin 143, 196, 222, 272, 278, 283, 290, Bromus inermis See brome, smooth 601, 661Ð662, 704, 724, 727, 748 Bromus marginatus See brome, mountain Ceanothus martinii See ceanothus, Martin Bromus riparius See brome, meadow ceanothus, prostrate 601, 659Ð661, 705, 748 Bromus tomentellus See brome, subalpine Ceanothus prostratus See ceanothus, prostrate

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-3 ceanothus, redstem 142Ð143, 222, 266Ð267, 272, clover, alsike 261, 264, 427, 464, 702 601, 662Ð665, 668, 704, 748 Aurora 464 Ceanothus sanguineus See ceanothus, redstem Dawn 464 ceanothus, snowbrush 196, 272, 283, 288, 290, 601, clover, Castle Valley See saltbush, Castle Valley clover 665Ð668, 705, 748 clover, red 464 Ceanothus velutinus See ceanothus, snowbrush Dollard 464 ceanothus, wedgeleaf 601, 656Ð657, 705, 749 Kenland 464 Celtis reticulata See hackberry, netleaf Lakeland 464 Ceratoides lanata See winterfat Pennscott 464 Ceratoides lanata subspinosa See winterfat, foothills clover, strawberry 261Ð262, 264, 427, 463Ð464, 702, Ceratoides latens See winterfat, Pamirian 724 Cercocarpus betuloides See mountain mahogany, clover, white 464 birchleaf coarse fragment content 41, 43 Cercocarpus intricatus See mountain mahogany, cold tolerance 128 littleleaf Coleogyne ramosissima See blackbrush Cercocarpus ledifolius See mountain mahogany, columbine, Colorado 259, 269, 427 curlleaf communities, plant 199 See also site preparation and Cercocarpus montanus See mountain mahogany, true seeding prescriptions; vegetation types certification, seed 734 companion species 62 certified seed See seed, certified compatibility chain See anchor chain perennial grasses 122 chaining 70, 228Ð229, 231, 233 seeding and species 122, 133 cheatgrass 102, 107, 254Ð255, 310 competition, plant See plant competition cheatgrass communities 9, 253Ð256, 294, 601 competitiveness of species 259, 261Ð262, 269, 271, checker-mallow, Oregon 264 273, 279Ð280, 282, 284Ð287, 289, 291Ð293 chemical plant control 62 composite shrubs 493 chenopod 467 container stock 739Ð742, 747, 754, 757, 764 cherry, Bessey 573Ð574, 601, 705, 749 copper 51Ð52, 54 cherry, bitter 217, 222, 272, 574Ð575, 601, 705, 749 Cornus sericeus See dogwood, redosier cherry, Nanking 577 Cornus stolonifera See dogwood, redosier cherry, western sand See cherry, Bessey Coronilla varia See crownvetch chokecherry 135, 142Ð143, 196, 266Ð267, 270, 272, Cotoneaster acutifolia See cotoneaster, Peking 278, 283, 288, 290, 577Ð579, 601, 705, 724, 730, cotoneaster, Peking 272, 278, 555Ð556, 705 749 cottonwood, Fremont 266Ð267 Chrysothamnus See rabbitbrush cottonwood, narrowleaf 266Ð267, 749 Chrysothamnus albidus See rabbitbrush, alkali cover See wildlife habitat, cover Chrysothamnus depressus See rabbitbrush, dwarf Cowania mexicana See cliffrose, Stansbury Chrysothamnus greenei See rabbitbrush, Greene’s Cowania stansburiana See cliffrose, Stansbury Chrysothamnus linifolius See rabbitbrush, spreading cowparsnip 216, 259, 264, 269, 426Ð427, 441Ð442, Chrysothamnus nauseosus See rabbitbrush, rubber 702, 724, 727 Chrysothamnus parryi See rabbitbrush, Parry Crataegus douglasii See hawthorn, Douglas Chrysothamnus vaseyi See rabbitbrush, Vasey crown divisions 762 Chrysothamnus viscidiflorus See rabbitbrush, low crownvetch 135, 196, 259, 264, 269, 271, 277, 282, chukar partridge 170 289, 426Ð427, 437Ð438, 702, 724, 731, 741 cinquefoil 749 Chemung 438 cinquefoil, bush 601, 705 See also cinquefoil, shrubby Emerald 438 cinquefoil, gland 259, 261, 264, 427 Penngift 438 cinquefoil, shrubby 260, 262, 266, 267, 270, 290, crude fat 177, 180 569Ð571 crude fiber 177, 180 Clematis ligusticifolia See virginsbower, western crude protein 177, 180 cliffrose, Stansbury 132, 135, 138, 142Ð143, 185, 196, cultivars 129, 138, 298, 734 272Ð273, 276, 278, 280, 283, 556Ð559, 562, 579, Cupressus arizonica See cypress, Arizona 601, 705, 725, 727, 730, 741, 749 curly grass See galleta climate 29, 34Ð35, 124 currant, golden 135, 143, 196, 266Ð267, 601, clover 463 690Ð691, 705, 725, 727, 741, 749

Index-4 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 currant, sticky 260, 602, 693Ð694, 705, 749 Elymus elymoides See squirreltail, bottlebrush currant, wax 260, 602, 691Ð693, 705, 749 Elymus giganteus See wildrye, mammoth cut stump treatment 92 Elymus glaucus See wildrye, blue cuttings, hardwood 761Ð762 Elymus junceus See wildrye, Russian cuttings, stem 739, 741 Elymus lanceolatus lanceolatus See wheatgrass, cypress, Arizona 276, 278, 602, 632Ð633, 705, 749 thickspike Elymus lanceolatus wawawai See wheatgrass, D bluebunch; wheatgrass, Snake River Dactylis glomerata See orchardgrass Elymus salinus See wildrye, Salina daisy, oxeye 427 Elymus simplex See wildrye, alkali debearder 713 Elymus trachycaulus See wheatgrass, slender deer 159, 161, 176, 515 Elymus triticoides See wildrye, creeping deervetch, birdfoot 427 Elymus wawawaiensis See wheatgrass, Snake River; depredation 144Ð145, 163, 768 wheatgrass, bluebunch Deschampsia caespitosa See hairgrass, tufted Elytrigia repens x Pseudoroegneria spicata See desert almond 572Ð573 wheatgrass, hybrid: NewHy desert blite 491 See also sumpweed, desert energy 176Ð177 dibbles 764Ð765 ephedra, green 132, 135, 142Ð143, 196, 272Ð273, digestibility 181 276, 278Ð280, 283Ð285, 602, 645Ð646, 705, 725, disease, plant 601 727, 730, 741, 749 disks 66Ð69 ephedra, Nevada 273, 276, 278Ð279, 281, 284Ð285, Distichlis spicata See saltgrass, inland 293, 602, 644Ð646, 705, 725, 727, 749 dixie sager 67, 69, 70 Ephedra nevadensis See ephedra, Nevada dogwood, creek See dogwood, redosier ephedra, Torrey 705, 749 dogwood, redosier 262, 266Ð267, 602, 630Ð632, 705, Ephedra viridis See ephedra, green 749 eriogonum, cushion 269, 271, 277, 427, 705 dozers and blades 84 Eriogonum heracleoides See eriogonum, Wyeth drags 71 eriogonum, sulfur 272, 283, 602, 652, 653Ð654, 749 drill seeding 205 Sierra sulfur buckwheat 654 drills 73 Eriogonum umbellatum See eriogonum, sulfur horizon 75 eriogonum, Wright 652, 654 no-till 75 Eriogonum wrightii See eriogonum, Wright rangeland 73Ð75, 228 eriogonum, Wyeth 143, 196, 276, 278, 283, 602, 652, Truax 75 652Ð653, 706, 727, 749 Tye 75 establishment 131 dropseed, sand 135, 142, 182Ð183, 185, 196, 262, esthetics 60 273, 279Ð280, 284, 291, 293, 308, 311, 414Ð416, F 700 drought tolerance 127Ð128 Fallugia paradoxa See Apache plume Dybvig separator 714 fences 172 fertilizers 53, 54, 55 E application 55Ð56 ecotypes 126, 200 elemental composition 54 edges, vegetative 111 fescue, Alpine See fescue, sheep elaeagnus, autumn 602, 705 fescue, Arizona 311 Elaeagnus commutata See silverberry Redondo 299 elderberry, blue 135, 143, 196, 270, 272, 276, 278, fescue, bluebunch See fescue, Idaho 283, 288, 290, 602, 624Ð625, 705, 741, 748 fescue, desert 700 elderberry, red 260, 266Ð267, 270, 290, 602, 625Ð627, fescue, hard See fescue, hard sheep 705 fescue, hard sheep 132, 135, 138, 142, 196, 269, 271, elk 111, 158Ð164, 176 273, 275, 277, 280, 282, 284, 286Ð287, 289, 294, Elymus angustus See wildrye, Altai 308, 311, 389, 700, 724, 728 Elymus canadensis See wildrye, Canada Durar 299, 390 Elymus cinereus See wildrye, Great Basin fescue, Idaho 102, 139, 182Ð184, 280, 282, 284, Elymus dahuricus See wildrye, Dahurian 286Ð287, 294, 308, 311, 386Ð388, 700

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-5 Joseph 299, 388 fireweed 264, 427 Nezpurs 299, 388 fixed-wing aircraft 78 fescue, meadow 308, 311, 731 flame thrower 85 fescue, red 132, 269 flax, Lewis 132, 135, 138, 142, 196, 264, 271, 273, fescue, reed 308, 384 See also fescue, tall 275, 277, 279Ð280, 282, 284, 286Ð287, 294, fescue, sheep 271, 273, 280, 282, 287, 289, 308, 311, 425Ð427, 443Ð444, 702, 727Ð728, 741 388, 700 Appar 444, 724 Covar 299, 390 fleabane, Bear River 259 MX86 299 fleabane, Oregon 259 sulcate 259, 271, 275, 277, 280, 282, 389 fluted shaft seeder 81 fescue, spike 308, 311 foliage application 92Ð93 fescue, tall 209, 259, 262, 264Ð265, 311, 384Ð386 forage production 60 Alta 269, 299, 311, 385 forbs 132, 135, 138, 142, 182Ð183, 185, 259, 702, Fawn 299, 385 724, 727, 731, 741, 763 varieties 386 forestiera, New Mexico 602, 706 fescue, Thurber 269, 308, 311 fox 178 Festuca arizonica See fescue, Arizona foxtail, creeping 259, 289, 308, 310, 346Ð349, 700 Festuca arundinacea See fescue, tall Garrison 300, 349 Festuca idahoensis See fescue, Idaho foxtail, meadow 206, 259, 261Ð262, 264Ð265, 269, Festuca ovina See fescue, sheep 289, 308, 310, 346Ð349, 700, 724 Festuca ovina duriuscula See fescue, hard sheep Dan 300, 349 filaree See alfileria foxtail, reed See foxtail, creeping fire effects 102Ð104, 107, 113Ð114 Fraxinus anomala See ash, singleleaf big sagebrush-cheatgrass 106 bitterbrush, antelope 103 G bluegrass 102 galleta 183Ð184, 250, 279, 293, 308, 311, 390Ð392 cheatgrass 102 Viva 300, 391 fescue, Idaho 102 geranium 132, 741 habitat management 113 geranium, Richardson 426Ð427, 438, 702, 724 horsebrush 103 Geranium richardsonii See geranium, Richardson intensity 113 geranium, sticky 259, 261, 269, 289, 426, 427, 439, junegrass, prairie 102 702 mountain mahogany, curlleaf 103 Geranium viscosissimum See geranium, sticky needlegrass 102 germination 601 oak, Gambel 103 giant hyssop, nettleleaf 427 plant succession 114 globemallow 132, 135, 196 rabbitbrush 103 globemallow, gooseberryleaf 142, 182, 273, 275, 279, reedgrass, plains 102 280, 284Ð285, 287, 292Ð293, 427, 461Ð462, 702, ricegrass, Indian 102 727, 741 riparian 112 globemallow, scarlet 273, 279, 284Ð286, 291, 293, sagebrush 103 427, 462, 702, 741 severity 113 ARS 2936 462 snowberry 103 globemallow, stream 427 squirreltail, bottlebrush 102 goat's beard See salsify, vegetable-oyster wheatgrass 102 goldeneye, Nevada 271, 273, 275, 280, 284, 427, 466, fire ignitor 85 703 aerial ignitors 85 goldeneye, showy 132, 135, 142, 196, 259, 269, 271, flame thrower 85 275, 277, 282, 287, 289, 427, 465, 466, 703, 724, helitorch 85 727, 731, 741 ping-pong ball injector 85 goldenrod, Canada 269, 271, 289, 427, 460Ð461, 703, teratorch 85 763 fire intensity 113 goldenrod, low 259, 269, 427 fire severity 113 goldenrod, Parry 277, 427 firebreak 114, 115 goldenweed 534, 706 fireline 115, 117 gouger 84 fireplow 84

Index-6 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 grama, black 308, 355Ð356 hammermill 713 Nogal 300, 356 Hansen seed dribbler 79 Sonora 356 Haplopappus See goldenweed grama, blue 308, 310, 356Ð358 Haplopappus bloomeri See rabbitbrush, goldenweed Hachita 300, 358 harvester Lovington 300, 358 sprigger 763 grama, sideoats 308, 310, 353Ð355 harvesting, seed 712 Butte 300 hawthorn, Columbia 561 El Reno 300 hawthorn, Douglas 266Ð267, 560Ð561, 602, 706, 750 Haskell 300 hawthorn, river See hawthorn, Douglas Killdeer 300, 354 heat damage 103 Niner 300, 354 Hedysarum boreale See sweetvetch, Utah Pierre 300, 355 helianthella, oneflower 182Ð183, 185, 259, 270, 277, Premier 300 426Ð427, 441, 703, 724 Trailway 300 Helianthella uniflora See helianthella, oneflower Vaughn 300, 355 helicopter 78Ð79 gravity separators 715 helitorch 85 Grayia brandegei See hopsage, spineless Heracleum lanatum See cowparsnip Grayia spinosa See hopsage, spiny herbicide effects 90Ð91, 96 grazing 2, 3, 128Ð129, 601 herbicide sprayer 85Ð87 plant control 59 herbicide types 94, 97 grazing management 194Ð198 acid equivalent 97 greasewood 470 nonselective 94 greasewood, Bailey 469, 490, 706 selective 94 greasewood, black 135, 195, 247, 262, 266, 267, 292, soil sterilant 94 469, 472, 490, 491, 602, 706, 725, 741, 750 toxicant 97 greasewood, black communities 9, 249Ð250, 292, herbicides 89, 95, 167, 255 308Ð309, 601 application 92, 98 Great Basin Experimental Range 16 broadcast ground application 93 Great Basin Station 3, 4, 16Ð17 broadcast spray application 92 greenhouse bioassay 53 cut stump treatment 92 grid ball 92 foilage 92 grinder-macerator 715 foilage spray 93 ground broadcast seeding 76, 205 grid ball 92 groundsel, butterweed 259, 264, 270Ð271, 289, 427, rate, calculation 97Ð98 459Ð460, 702 safety considerations 99 grouse 178Ð179 soil application 92 blue 170 soil injection 92 Columbian sharp-tail 168Ð169 soil surface placement 92 ruffed 169Ð171 stem 92 sage See sage-grouse timing of application 98 growth rate 128 trunk base spray 92 Gutierrezia See matchbrush; snakeweed trunk injection 92 Gutierrezia microcephala See matchbrush, small wipe on 92, 93 headed common name Gutierrezia sarothrae See snakeweed, broom clopyralid 95 dicamba 95 H glyphosate 95 habitat improvement projects 156 Oust 95 hackberry, netleaf 602, 696Ð698 paraquat 95 hairgrass, tufted 259, 261, 264Ð265, 289, 308, 310, picloram 95 369Ð371 tebuthiuron 95 Norcoast 301, 371 triclopyr 95 Nortran 301, 371 type 95 Peru Creek 301, 371 uses 90

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-7 Hesperostipa comata See needle-and-thread Juniperus osteosperma See juniper, Utah Hilaria jamesii See galleta Juniperus scopulorum See juniper, Rocky Mountain hoedad 764 Holodiscus discolor See oceanspray, creambush K Holodiscus dumosus See spiraea, rock kinnikinnik See manzanita, bearberry honeylocust 602, 706 Kochia americana See molly, gray honeysuckle 731 kochia, Belvedere 703 honeysuckle, bearberry 602, 623Ð624 kochia, forage 135, 138, 142, 196, 273, 276, 278, 279, honeysuckle, orange 602, 623 281, 283, 284, 285, 292, 293, 294, 469, 472, 489, honeysuckle, Tatarian 262, 266Ð267, 272, 706, 750 490, 603, 707, 730, 741 honeysuckle, Utah 262, 272, 602, 623Ð624, 706, 750 Immigrant 490, 725 honeysuckle, western trumpet See honeysuckle, Kochia prostrata See kochia, forage orange Koch’s postulates 188 hopsage, spineless 273, 291, 293, 469, 472, 487, 488, Koeleria cristata See junegrass, prairie 602, 706, 727 Koeleria macrantha See junegrass, prairie hopsage, spiny 279, 281, 284, 291, 293, 469, 487, Koeleria nitida See junegrass, prairie 488, 602, 706, 725, 750 Horizon drill 75 L horsebrush 103, 534Ð536 horsebrush, cottonthorn 706 land imprinter 67, 72 horsebrush, gray 495, 535, 706 larkspur, tall 204 horsebrush, littleleaf 495, 706 layering 762 horsebrush, Nuttall 495, 706 crown divisions 762 horsebrush, spiny 495 runners and stolons 762 horses 176, 178, 179 shoots 762 human activities 194, 198 stem layers 762 hydroseeder 82 suckers 762 layers 762 I legumes 136, 137 Lepidospartum latisquamatum See scalebroom Indian apple 272, 276, 278, 283, 566Ð567, 604, 711, Leymus angustus See wildrye, Altai 727, 731, 741 Leymus cinereus See wildrye, Great Basin inkbush See iodine bush Leymus racemosus See wildrye, mammoth Intermountain Forest and Range Experiment Station 3, Leymus salinus See wildrye, Salina 16 See also Intermountain Research Station Leymus simplex See wildrye, alkali Intermountain Research Station 4, 5, 17 Leymus triticoides See wildrye, creeping interseeder 81 ligusticum, Porter 259, 270, 426Ð427, 442Ð443, 703, fluted shaft seeder 81 727 Truax single row seeder 81 Ligusticum porteri See ligusticum, Porter interseeding 142, 742Ð744 lilac, common 707, 750 iodine bush 469, 491 Linum perenne lewisii See flax, Lewis iris, German 427, 741 livestock grazing See grazing; grazing management iron 51, 52, 54 locust, black 603, 707, 750 J Lomatium kingii See lomatium, Nuttall lomatium, narrowleaf See lomatium, nineleaf jointfir See ephedra, green; ephedra, Nevada lomatium, nineleaf 426, 428, 444Ð445, 703, 724, 728 junegrass, prairie 102, 271, 275, 277, 282, 287, 308, lomatium, Nuttall 259, 270Ð271, 277, 426, 428, 311, 392Ð393, 700 445Ð446, 703, 727 juniper 108 Lomatium triternatum See lomatium, nineleaf juniper, creeping 602, 633Ð635 Lonicera ciliosa See honeysuckle, orange juniper, Rocky Mountain 602, 638Ð640, 706, 750 Lonicera involucrata See honeysuckle, bearberry juniper, Utah 182Ð183, 185, 602, 635Ð638, 638, 707, Lonicera utahensis See honeysuckle, Utah 750 lupine 135, 196, 741 juniper woodlands 38 lupine, mountain 142, 259, 270, 282, 289, 703, 724, juniper-pinyon See pinyon-juniper 727, 728 Juniperus horizontalis See juniper, creeping lupine, Nevada 271, 277, 280, 284, 428

Index-8 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 lupine, silky 260, 270Ð271, 277, 282, 287, 289, 426, mockorange, littleleaf 689 428, 446Ð447, 703, 724, 727Ð728 moldboard plow 66 lupine, silvery 428, 448, 703 molly, gray 469, 488, 603 Lupinus argenteus See lupine, silvery molybdenum 51Ð52, 54 Lupinus sericeus See lupine, silky moose Shiras 164 M moth, sagebrush defoliator 514 MacLeod 764, 765 mountain ash, American 707, 750 magnesium 51Ð52, 54 mountain ash, Greene’s 266Ð267, 270, 272, 592Ð594, Mahonia aquifolium See barberry, shining 601, 603 mahonia, creeping See Oregon grape mountain ash, Sitka 594 Mahonia fremontii See barberry, Fremont mountain brush communities 9, 195, 216, 217, 219, Mahonia repens See Oregon grape 271, 308Ð309, 601 management 28, 32 mountain lover 222, 266Ð267, 603, 707 considerations 26 mountain mahogany, birchleaf 138, 544Ð545, 707, 750 climate 29 mountain mahogany, curlleaf 103, 132, 135, 142Ð143, impacts on adjacent areas 31 182, 183, 185, 196, 217, 272, 276, 278, 283, resource values 31 545Ð550, 603, 707, 725, 727, 730, 741, 750 soil condition 27Ð28, 30Ð31 mountain mahogany, Intermountain 546 strategy 28 mountain mahogany, Intermountain curlleaf 549 vegetation condition 26Ð27 mountain mahogany, littleleaf 274, 276, 545, 549, 603, manganese 51Ð52, 54 707, 751 manzanita, bearberry 601, 647Ð648, 707, 750 mountain mahogany, true 132, 135, 138, 142Ð143, maple 195 185, 196, 272, 276, 278, 283, 549, 550Ð552, 603, maple, bigtooth 270, 603, 750 707, 725, 727, 730, 741, 751 maple, Rocky Mountain 216, 272, 278, 600Ð608, 707, Montane 552 750 muhly, spike 731 matchbrush 493, 534 El Vado 301 matchbrush, small head 495 mule deer 158, 163, 178 McSweeney-McNary Forest Research Act of 1928 3 ranges 162 meadowrue, Fendler 260 multiple use 131 mechanical plant control 62 muttongrass 273, 275, 282, 286Ð287 See also blue- brushland plow 66 grass, mutton disk plow 66 muttongrass, longtongue 311 disk-chain 66 N moldboard plow 66 off-set disk 66 Nassella viridula See needlegrass, green personnel 78 natural spread 134 Medicago falcata See alfalfa, sicklepod needle-and-thread 182Ð184, 220, 228, 237, 243, 256, Medicago sativa See alfalfa 273, 275, 277, 279, 280, 282, 284Ð286, 293Ð294, medick, black 261Ð262, 264, 270, 428 308, 311, 418Ð419, 700, 731 medusahead 254Ð255 needlegrass 102, 139 medusahead communities 253Ð256, 294 needlegrass, Columbia See needlegrass, subalpine Melilotus officinalis See sweetclover, yellow needlegrass, green 135, 142, 269, 271, 275, 277, 282, Mertensia arizonica See bluebell, tall 287, 289, 308, 311, 422Ð424, 700 milkvetch, Canadian 703 Cucharas 424 milkvetch, cicer 132, 135, 138, 142, 196, 261, 270, Lodorm 301, 424 271, 277, 282, 289, 425Ð426, 428, 433Ð434, 703, needlegrass, Letterman 269, 282, 289, 308, 311, 724, 728, 731, 741, 763 419Ð421, 700 Lutana 434 needlegrass, subalpine 259, 269, 308, 311, 416Ð418, milkvetch, Snake River 428 420 milkvetch, tall 428 varieties 418 mixed seedings 130Ð134 needlegrass, Thurber 280, 284, 287, 294, 308, 311, mockorange, Lewis 687Ð689, 707 421Ð422 Waterton 689 ninebark, dwarf 568

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-9 ninebark, mallow 266Ð267, 567Ð569, 603, 707, 751 peavine, Utah 270, 428 ninebark, mountain 568 penstemon, bush 272, 603, 694Ð696, 707, 751 ninebark, Pacific 568 Penstemon cyananthus See penstemon, Wasatch nitrogen 51, 52, 54 penstemon, Eaton 139, 271, 277, 280, 282, 287, 289, nitrogen-free extract 177, 180 428, 456, 703, 731 no-till drill 75 Richfield 457 nurseries and facilities 747 Penstemon eatonii See penstemon, Eaton nutrient content 112 penstemon, firecracker 456 nutrient requirements of range animals 176 Penstemon fruticosus See penstemon, bush calcium 178 Penstemon humilis See penstemon, low dry material 176 penstemon, littlecup 428 minerals 178 penstemon, low 206, 271, 280, 282, 289, 428, 457, phosphorus 178 703, 724, 763 protein 177, 178 Penstemon pachyphyllus See penstemon, thickleaf vitamins 179 penstemon, Palmer 132, 135, 138, 142, 196, 271, 273, nutrient values of range plants 180Ð184 275, 277, 279Ð280, 282, 284Ð287, 293Ð294, 425, 428, 454Ð456, 703, 727Ð728, 731 O Cedar 456 oak, Gambel 103, 135, 183, 185, 195, 217, 218, 603, Penstemon palmeri See penstemon, Palmer 648Ð650, 707, 741, 751 penstemon, Rocky Mountain 132, 135, 260, 270Ð271, oatgrass, tall 135, 206, 259, 261, 269, 271, 282, 287, 277, 283, 289, 425, 428, 457 289, 308, 310, 351Ð353, 701 penstemon, Rydberg 260, 270, 428, 457 Tualatin 353, 354 Penstemon rydbergii See penstemon, Rydberg oceanspray, bush 565 See also oceanspray, spiraea penstemon, sidehill 428 oceanspray, creambush 563Ð565, 603 Penstemon strictus See penstemon, Rocky Mountain Office of Grazing Studies 3 penstemon, thickleaf 277, 428, 457, 703, 731 olive, autumn 751 penstemon, toadflax 277, 428 oniongrass 259, 308, 311, 701 penstemon, Wasatch 260, 270Ð271, 277, 283, 289, Onobrychis viciaefolia See sainfoin 428, 456, 703 orchardgrass 132, 135, 142, 196, 205, 206, 216, 220, Peraphyllum ramosissimum See Indian apple 255, 259, 264Ð265, 269, 271, 273, 275, 277, 280, persistence 282, 287, 289, 308, 310, 367Ð369, 701, 724, 728 species 129 Berber 368 personnel 63 Latar 301, 368 Phalaris arundinacea See canarygrass, reed Paiute 138, 301, 368, 724, 728 pheasant 159, 178Ð179 Potomac 301 Philadelphus lewisii See mockorange, Lewis Oregon grape 270, 614Ð616, 731 See also barberry, Phleum alpinum See timothy, alpine creeping Phleum pratense See timothy Oryzopsis hymenoides See ricegrass, Indian phosphate, single super 54 Osmorhiza occidentalis See sweetanise phosphate, trebel super 54 phosphoric acid (liquid) 54 P phosphorus 51, 52, 54, 178, 179 Physocarpus malvaceus See ninebark, mallow painted-cup 428 pine, ponderosa 195 painted-cup, sulphur 260 pine, ponderosa communities 222Ð223, 309, 601 Pascopyrum smithii See wheatgrass, western ping-pong ball injector 85 pathogen-free planting stock 189 pinyon 108 pathogens, plant 187Ð190 pinyon-juniper 195, 224, 225, 279 peachbrush, Anderson 707, 751 See almond, desert pinyon-juniper communities 9, 37, 223Ð224, 227Ð228, peachbrush, desert 274, 276, 284, 291, 293, 575Ð577, 273, 279, 308, 601, 638 603, 707, 725 pipe harrow 67, 71 pearly everlasting 763 plant communities, diverse 130 peashrub, Siberian 731, 751 plant community indicators 124 peavine, flat 428 plant competition 21, 57, 61, 112, 134, 202, 207Ð208, peavine, perennial 428 211, 214, 218, 221, 226, 235, 238, 239Ð240, 243, peavine, thickleaf 270, 428 245, 247, 249, 251Ð252, 255, 257Ð258

Index-10 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 controlling 57, 65, 147Ð148 Prunus andersonii See almond, desert biological 58, 59 Prunus besseyi See cherry, Bessey burning 60 Prunus emarginata See cherry, bitter chemical 62 Prunus fasciculata See peachbrush, desert fallowing 62 Prunus spinosa See blackthorn grazing 59 Prunus tomentosa See cherry, Nanking herbicides 58 Prunus virginiana See chokecherry logging 60 Psathyrostachys juncea See wildrye, Russian mechanical 58, 62, 65 Pseudoroegneria spicata See wheatgrass, bluebunch objectives 60 Pseudoroegneria spicata inerme See wheatgrass, species 21 beardless successional changes 59Ð60 pure live-seed (PLS) 22, 139, 736 plant control Purshia glandulosa See bitterbrush, desert herbicides 89 Purshia tridentata See bitterbrush, antelope plant disease fungal diseases 516 Q plant tissue analysis 53 quackgrass 310 planting bar 764Ð767 quail 178, 179 planting hoe 764Ð765 quailbush See saltbush, quailbush planting mixtures, advantages 130Ð133 Quercus gambelii See oak, Gambel planting season 23, 150Ð151 planting shovels 764Ð765 R planting stock 745Ð746 plows rabbit 178 brushland 66Ð67 rabbitbrush 103, 493Ð496, 522Ð524 disk 66 rabbitbrush, alkali 495, 533Ð534, 708 moldboard 66 rabbitbrush, Douglas 603 root 67, 73 rabbitbrush, dwarf 495, 524, 603, 708 PLS See pure live-seed (PLS) rabbitbrush, goldenweed 534 plum, American 262, 571Ð572, 603, 708, 751 rabbitbrush, Greene’s 495, 533Ð534, 708 plume, Apache See Apache plume rabbitbrush, low 135, 182Ð183, 185, 196, 260, 279, Poa alpina See bluegrass, alpine 285Ð286, 291, 293Ð294, 495, 531Ð533, 603, 708 Poa ampla See bluegrass, big hairy low 533 Poa canbyi See bluegrass, Sandberg mountain low 142, 288, 290, 532, 708, 725, 741 Poa compressa See bluegrass, Canada narrowleaf low 533, 708 Poa fendleriana See bluegrass, mutton stickyleaf low 532Ð533, 708 Poa longiligula See bluegrass, mutton rabbitbrush, Nevada 530 Poa nevadensis See bluegrass, Sandberg rabbitbrush, Parry 260, 272, 495, 529Ð531, 603, 708 Poa pratensis See bluegrass, Kentucky rabbitbrush, rubber 132, 135, 182Ð183, 185, 196, 251, Poa sandbergii See bluegrass, Sandberg 262, 495, 525Ð529, 603, 751 Poa scabrella See bluegrass, Sandberg green rubber 529, 708 Poa secunda See bluegrass, big; bluegrass, Sandberg green-stemmed rubber 526, 741 ponderosa pine communities 220, 271, 308 leafless rubber 708 poplar 751 leiospermus rubber 708 postfire management 119 mountain rubber 270, 272, 276, 278, 283, 288, 529, potash 54 708, 725 potassium 51Ð52, 54 mountain whitestem rubber 708 Potentilla fruticosa See cinquefoil, shrubby threadleaf rubber 266Ð267, 528Ð529, 708, 741 precipitation 21, 34Ð35 tubinatus rubber 708 average annual 34 whitestem mountain and basin rubber 270, 272, pronghorn antelope 111, 164Ð165 274, 276, 278, 281, 283Ð284, 286, 288 propagation See seed propagation; vegetative whitestem rubber 138, 142Ð143, 525Ð526, 528, 725, propagation 727, 730, 741 protein 177 rabbitbrush, small 603 proximal analysis 180 rabbitbrush, spreading 291, 495, 524, 603, 708, 741 Prunus americana See plum, American rabbitbrush, Vasey 495, 531, 709

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-11 rails 71Ð72 Ribes viscosissimum See currant, sticky range, season of use 182Ð184 ricegrass, Indian 102, 132, 135, 138, 142, 182, 184, Range Seeding Equipment Committee 17 196, 273, 275, 277, 279, 280, 282, 284, 285, raspberry, black 590, 601 286Ð287, 291, 293Ð294, 308, 311, 393Ð396, 701, redtop See bentgrass, redtop 731 reedgrass, bluejoint 264Ð265, 310 Nezpar 301, 396 Sourdough 301 Paloma 301, 397 reedgrass, chee 259, 264Ð265, 310, 701, 763 Ribstone 397 reedgrass, plains 102 Rimrock 301, 397 rehabilitation 200 See also site preparation and riparian communities 9, 209Ð210, 212Ð213, 263Ð267, seeding prescriptions 309, 601, 668 resistance, host 190 roads 172, 198 resource values 31 rockspirea 565Ð566, 709 restoration 19, 200 See also revegetation; site prepa- Rocky Mountain Forest and Range Experiment Station ration and seeding prescriptions 3 See also Rocky Mountain Research Station history 1, 15Ð17 Rocky Mountain Research Station 5 human activities 198 rodents 144 restored sites roller chopper 83 grazing 194 root cuttings 762 human activities 194 root-plow 67, 73 management 193 Rosa woodsii ultramontana See rose, Woods post-treatment 194 rosaceous shrubs 539 revegetation 19, 29, 200, 470, 717, 746 See also rose, Woods 143, 260, 262, 266Ð267, 270, 272, 278, restoration; site preparation and seeding 283, 288, 586Ð590, 603, 709, 741, 751 prescriptions Roundup 96 artificial 29 row spacing 153 considerations 19 Rubus leucodermis See raspberry, black cost 32 Rubus parviflorus See thimbleberry impacts on adjacent areas 31Ð32 runners and stolons 762 plant competition 21 rush, Baltic 263 plant material availability 32 rush, Drummond 263 planting season 23 rush, longstyle 263 precipitation 21 rush, swordleaf 263 pure live seed (PLS) 22 rush, Torrey 263 resource values 31 Russian olive 709 seed mixture 22 rye, mountain 132, 135, 142, 196, 269, 273, 275, 277, seedbed 23 280, 282, 284, 286Ð287, 294, 309, 311, 724 site-adapted species and populations 21 rye, winter 273, 275, 277, 727 soil conditions 27Ð28, 30 ryegrass, Italian 311 soil types 20 ryegrass, meadow 311 terrain 20 ryegrass, perennial 264, 265, 311 vegetation condition 26Ð27 weeds 31 S history 15Ð17 sacaton, alkali 196, 262, 264Ð265, 279, 286, 309, 311, human activities 198 413Ð414, 701 impact on resources 28 Salado 301, 414 range 19 Saltalk 301, 414 site suitability 29Ð30 sage, birdsfoot 495, 522 wildland 19 sage, fringed 274, 495, 505Ð506, 604, 709, 725 Rhizobium 146 sage, longleaf 495, 522, 709 rhizomes 763 sage, Louisiana 260Ð261, 264, 270Ð271, 278, 289, Rhus aromatica See sumac, skunkbush 426, 428, 430Ð431, 703, 741, 763 Rhus glabra See sumac, Rocky Mountain smooth Summit 431 Ribes aureum See currant, golden sage, oldman See sandsage Ribes cereum See currant, wax

Index-12 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 sage, purple 709, 751 sagebrush, low 206, 232, 279, 281, 283, 285, 495, sage, sand See sandsage 499Ð500, 506, 604, 709 sage, tarragon 278, 428 low communities 244, 286 sage-grouse 159, 166Ð168 sagebrush, pygmy 495, 509Ð510, 604, 709 sagebrush 108, 236, 493, 494Ð496 sagebrush, Rothrock 520, 522 sagebrush communities 115, 234Ð237 See also site sagebrush, sand 293, 709 See also sandsage preparation and seeding prescriptions sagebrush, scabland See sagebrush, stiff sagebrush, alkali 495, 506Ð507 sagebrush, silver 232, 260, 266Ð267, 272, 283, 290, sagebrush, alkali communities 245 495, 502Ð503, 604, 709, 741 sagebrush, big 132, 181Ð183, 185, 196, 232, 294, Bolander silver 503 495, 514Ð521, 709, 751 mountain silver 502, 503 big communities 9 mountail silver communities 244 basin big 135, 138, 142Ð143, 195, 238, 266Ð267, plains silver 503 274, 276, 279, 281, 288, 292, 514Ð521, 604, 709, silver communities 289 725, 727, 730, 741 sagebrush, stiff 495, 510Ð511, 604, 709 Hobble Creek 518 sagebrush, tarragon See sage, tarragon basin big communities 135, 238, 279, 280, 308Ð309, sagebrush, threetip 232, 288, 495, 521Ð522, 604 601 tall threetip 266Ð267, 522, 709 foothills big 272, 274, 276, 278, 279, 281, 283Ð284 threetip communities 244, 287 See also sagebrush, big: xeric big Wyoming threetip 522 mountain big 135, 138Ð139, 142, 195, 232, 260, sainfoin 132, 135, 138, 196, 261, 270, 272, 275, 278, 266Ð267, 270, 272, 276, 278, 283, 288, 283, 289, 453, 703, 724, 741 514Ð521, 604, 709, 725, 730, 741 Eski 454 Hobble Creek 518 Melrose 454 mountain big communities 239Ð240, 282, 308Ð309, Nova 454 601 Onar 454 spicate big 503, 514 See also sagebrush, big: Remont 454 subalpine big Runemex 454 subalpine big 206, 290 sainfoin, common 142, 428 subalpine big communities 289, 308Ð309 Salix See willow timberline big 244, 260, 289, 290, 520, 709 See also Salix bebbiana See willow, Bebb sagebrush, big: spicate big; sagebrush, big: Salix boothii See willow, Booth subalpine big Salix drummondiana See willow, Drummond timberline big communities 289 Salix exigua See willow, coyote Wyoming big 135, 138, 142, 195, 250, 251, 274, Salix geyeriana See willow, Geyer 279, 281, 286, 288, 291, 514Ð521, 604, 725, 730, Salix glauca See willow, grayleaf 741 Salix lasiandra caudata See willow, whiplash Gordon Creek 519 Salix lasiolepis See willow, arroyo Wyoming big communities 240Ð242, 279, 284, Salix lutea See willow, yellow 308Ð309, 601 Salix planifolia See willow, plainleaf xeric big 514, 517 See also sagebrush, big: foothills Salix scouleriana See willow, Scouler big Salix wolfii See willow, Wolf sagebrush, Bigelow 495, 500Ð501, 604, 709 salsify, vegetable-oyster 428, 703, 724, 727, 741 sagebrush, black 132, 142, 143, 185, 195, 196, 228, salt desert shrub communities 246 See also site 242, 274, 276, 285, 286, 291, 293, 495, 508Ð509, preparation and seeding prescriptions 604, 709, 725, 727, 730, 741, 751 saltbush 469 black communities 243, 285 saltbush, allscale 469, 482Ð483 sagebrush, budsage 604, 709 See also budsage saltbush, Australian 469 budsage communities 245 saltbush, big 469, 710 sagebrush, coaltown 503, 522 saltbush, Bonneville 469, 710 sagebrush, early 604 See sagebrush, alkali saltbush, broadscale 469, 472, 482Ð483, 710 sagebrush, fringed See sage, fringed saltbush, Castle Valley clover 292, 469, 472, 480Ð481, sagebrush, hotsprings 500 604, 710 sagebrush, longleaf See sage, longleaf saltbush, cattle 469, 710 sagebrush, Louisiana See sage, Louisiana saltbush, cuneate See saltbush, Castle Valley clover

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-13 saltbush, desert holly 469, 482Ð483, 710 seed saltbush, falcate 469, 710 afterripening 700, 726 saltbush, fourwing 129, 132, 135, 138, 142, 143, 182, certification 139, 719 183, 185, 196, 251, 262, 266, 267, 274, 276, 278, certified 137Ð138 279, 281, 283, 284, 286, 291, 292, 293, 469, 471Ð cleaning 699, 713Ð715 476, 604, 710, 725, 727, 730, 741, 751 collecting 699, 712, 718, 754 fourwing communities 251Ð252, 279 dormancy 127, 726, 730 Marana 475, 476 field 721, 754 Rincon 475, 476 germination 139, 205, 723, 724, 727Ð728, 730Ð732, Wytana 475, 476 746 saltbush, Gardner 209, 251, 262, 266Ð267, 279, 291Ð292, inoculation 146 469Ð470, 481Ð482, 604, 710, 725, 752 longevity 731 saltbush, Gardner complex 470, 481Ð482 maturity 700 saltbush, Garrett 710 noncertified 138 saltbush, mat 291, 469, 472, 479Ð480, 604, 710 orchard 718Ð719, 722, 754 saltbush, Navajo 469, 710 pathogens 146Ð147 saltbush, quailbush 469, 482Ð483, 604 pelleting 76, 146 Casa 483 pretreatment 145Ð146, 755 saltbush, robust 469 production 719 saltbush, shadscale 308, 604, 710, 752 propagation 747Ð748 shadscale communities 9, 248, 309, 601 purity 139, 700 saltbush, trident 469, 472, 710 quality 137 saltbush, wingless 469 storage 700, 715, 731, 754Ð755 saltgrass 199, 208Ð209, 212, 262, 264Ð265 stratification 700 saltgrass, desert 763 seed certification 733 See also certification, seed saltgrass, inland 195, 310, 371Ð373 seed, certified 22 inland communities 9, 208Ð209, 262, 308-309, 601 seed dribbler 79Ð80 Sambucus cerulea See elderberry, blue seed harvesters Sambucus racemosa See elderberry, red See elder- vacuum-type 713 berry, red seed mixture 22 sand sage See sandsage seed, noncertified 138 sandsage 495, 504Ð505, 604 seed production 601 Sanguisorba minor See burnet, small seed testing Sarcobatus vermiculatus See greasewood, black requirements 733 scalebroom 534, 537 seedbed 23, 126, 134 scalper 84 conditions 63 sedge, analogue 263 firmness 149, 150 sedge, beaked 263 preparation 121 sedge, black alpine 263 climatic conditions 124 sedge, blackroot 263 plant community indicators 124 sedge, Douglas 263 soil sedge, downy 263 divalent ions 125 sedge, golden 263 monovalent cations 125 sedge, Hepburn 263 seedbed preparation equipment 65 See also anchor sedge, hood 263 chains; cables; disks; plows sedge, Kellogg 263 drags 71 sedge, Nebraska 263 herbicide 91 sedge, ovalhead 259, 261, 289 land imprinter 67, 72 sedge, rock 263 pipe harrow 67, 71 sedge, russet 263 rails 71 sedge, slim 263 seeders 65 See also drills sedge, smallwing 263 aerial broadcasting 77Ð78 sedge, soft-leaved 263 fixed-wing aircraft 78 sedge, valley 263 helicopters 78Ð79 sedge, woolly 263 Brillion seeder 80 sedges 208, 263 broadcast 72, 76Ð77

Index-14 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 hand broadcasting 76 shadscale 132, 135, 143, 195, 196, 228, 243, 245Ð247, Hansen seed dribbler 79 248, 250Ð253, 257Ð258, 279, 291Ð292, 294, 469, hydroseeder 82 476Ð478 interseeder sheep 176, 178Ð179, 515 fluted shaft seeder 81 Shepherdia argentea See buffaloberry, silver Truax single row seeder 81 Shepherdia canadensis See buffaloberry, russet land imprinter 72 Shepherdia rotundifolia See buffaloberry, roundleaf seed dribbler 79Ð80 Shiras moose 164 surface seeder 80 shoots 762 surface seeders 80 shrub enhancement 235 thimble seeder 79Ð80 shrub-herb associations 134 Truax Seed Slinger 76 Shrubland Biology and Restoration Project 4 seeding 216 shrubs 132, 135, 138, 142, 182Ð183, 185, 260, 597, broadcast 141 704 depth 76, 152Ð153 silverberry 266Ð267, 604, 640Ð641 ecotype 124 Sitanion hystrix See squirreltail, bottlebrush mixed 201 site access 61 mixed seedings 130 site improvement 122 mixes 122, 129 site preparation and seeding prescriptions forbs 140 aspen communities 215, 309 native species 126 aspen openings 204Ð205 plant competition 21 aspen-conifer communities 9, 216, 601 planting season 23 Bailey's (1978) 8 precipitation 21 blackbrush communities 9, 252Ð253, 293, 308, 601 pure live seed (PLS) 22 brome, red communities 9, 253Ð256, 294 rate 22, 78 cheatgrass communities 9, 253Ð256, 294, 601 seed mixture 22 greasewood, black communities 9, 249Ð250, 309, seedbed 23 601 site-adapted species and populations 21 juniper woodlands 38 soil types 20 medusahead communities 253Ð256, 294 seeding practices 121 mountain brush communities 9, 219, 309, 601 rates 137, 139Ð141, 145 pine, ponderosa communities 222Ð223, 309, 601 requirements 135, 143 pinyon-juniper communities 9, 37, 223Ð224, 227Ð228, separate row seeding 154 273, 279, 308, 601, 638 single species seeding 130, 133 riparian communities 9, 209Ð210, 212Ð213, 263Ð267, species compatibility 133, 135 309, 601, 668 spot 142 sagebrush, alkali communities 245 seeding rates 259, 261, 269, 271, 273, 279Ð280, sagebrush, basin big communities 135, 238, 279, 284Ð287, 289, 291Ð293 280, 308Ð309, 601 seeding requirements sagebrush, big communities 9 row spacing 153 sagebrush, mountain big communities 239Ð240, spot seeding 142 282, 308Ð309, 601 seedling sagebrush, subalpine big communities 289, 308Ð309 establishment 133, 134 sagebrush, timberline big communities 289 growth rate 135 sagebrush, Wyoming big communities 240Ð242, vigor 135 279, 284, 308Ð309, 601 selenium 52 sagebrush, black communities 243, 285 Senecio serra See groundsel, butterweed sagebrush, budsage communities 245 serviceberry, dwarf Saskatoon 541 sagebrush communities 235Ð237 serviceberry, Saskatoon 132, 135, 142, 143, 196, 266, sagebrush, low communities 244, 286 267, 270, 272, 276, 278, 283, 288, 541Ð543, 604, sagebrush, silver communities 289 710, 725, 727, 741, 752 sagebrush, mountain silver communities 244 serviceberry, Utah 272, 274, 276, 541, 543Ð544, 604, sagebrush, threetip communities 244, 287 710, 725, 727, 752 saltbush, fourwing communities 251Ð252, 279

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-15 saltbush, shadscale 246Ð247, 291 reclamation potential 40 saltbush, shadscale communities 9, 248, 309, 601 aspect 47 saltgrass, inland communities 9, 208, 209, 262, 308, parent material 47 309, 601 texture 152 subalpine communities 601 soil application 92 subalpine herblands 9, 204, 205 soil carbon 44 weeds, annual communities 308, 601 soil conditions 36, 39Ð53, 41 weeds, annual lowland communities 9, 257, 258 bulk density 41, 42 wet and semiwet meadows 9, 168, 207, 208, 308, carbon 44 601 coarse fragment content 41, 43 winterfat communities 251 depth to limiting layer 41, 44 site-adapted species and populations 21 exchangeable sodium 41, 47 skunk cabbage 204 organic matter 41, 44 snakeweed 534, 536 permeability 41, 43 snakeweed, broom 495, 536 pH 41, 46 snowberry 103 salinity 41, 45, 46 snowberry, common 266Ð267, 604, 627, 710, 752 slope 41, 44 snowberry, desert 627Ð628 structure 41Ð42 snowberry, longflower 278, 604, 710, 752 texture 41Ð42 snowberry, longleaf 272 water-holding capacity 41, 43 snowberry, mountain 143, 206, 260, 266, 267, 270, soil factors See soil conditions 272, 276, 278, 283, 288, 290, 604, 629Ð630, 710, soil injection 92 727, 741, 752 soil reaction See soil, pH snowberry, western 266, 267, 604, 628Ð629 soil surface placement 92 snowberry, white See snowberry, common soil types 9, 20 snowmold 517 Solidago canadensis See goldenrod, Canada snowmold disease 516 Solomon plume, fat 260, 428 soapberry See buffaloberry, russet Solomon-seal, western 264 softwood cuttings 761Ð762 Sorbus scopulina See mountain ash, Greene’s soil 40, 125Ð126 Sorbus sitchensis See mountain ash, Sitka carbon 44 species selection condition 27Ð28, 30 climatic conditions 124Ð125 conditions 125 cold tolerance 128 erosion 28, 40 compatibility 122, 133 fertility 40, 52 cultivars 129, 137 fertilization 48 drought tolerance 127Ð128 improvement 40 growth rate 135 moisture 148Ð149 native species 126Ð128 nitrogen 136 palatability 128 nutrient 40, 47, 48, 49, 50 persistence 129 boron 51Ð52 seed dormancy 127 calcium 51 soils 125Ð126 concentrations 51 Sphaeralcea coccinea See globemallow, scarlet copper 51Ð52 Sphaeralcea grossulariifolia See globemallow, iron 51Ð52 gooseberryleaf magnesium 51Ð52 spikerush, common 263 manganese 51Ð52 spiraea 594 molybdenum 51Ð52 Spiraea betulifolia See spiraea, bridal wreath nitrogen 50Ð52 spiraea, birchleaf See spiraea, bridal wreath phosphorus 50Ð52 spiraea, bridal wreath 594Ð595, 604 potassium 51Ð52 Spiraea densiflora See spiraea, subalpine selenium 52 spiraea, Douglas 596, 711, 752 sulfur 50Ð51 Spiraea douglasii See spiraea, Douglas sulphate-sulphur 52 spiraea, subalpine 596 zinc 51Ð52 spiraea, white 594, 595 pH 45 spirea, rock 266Ð267, 288, 751 See also rockspirea

Index-16 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 spirea, rock spray 565 sweetvetch, Utah 132, 135, 142, 196, 272, 275, 278, Sporobolus airoides See sacaton, alkali 280, 283, 287, 426, 429, 439Ð440, 704, 724, Sporobolus cryptandrus See dropseed, sand 727Ð728, 741, 763 sprigger 763 Timp 440 squirrels 178 Symphoricarpos albus See snowberry, common squirreltail See squirreltail, bottlebrush Symphoricarpos longiflorus See snowberry, desert squirreltail, big 411 Symphoricarpos occidentalis See snowberry, western Sand Hollow 301, 412 Symphoricarpos oreophilus See snowberry, mountain squirreltail, bottlebrush 102, 135, 182Ð184, 196, 264, syringa See mockorange, Lewis 265, 271, 273, 275, 277, 279, 280, 282, 284Ð287, syringa, Lewis See mockorange, Lewis 291Ð294, 309, 311, 410Ð412, 701, 724, 728 Fish Creek 412 T Toe Jam Creek 412 tarweed, cluster 203 State Agricultural Experiment Stations 3, 4 temperature 35 steep-slope scarifier and seeder 87 tephritid flies 526 stem application 92 teratorch 85 stem cuttings 760 terrain 20Ð21, 36, 37 stem layers 762 Tetradymia See horsebrush Stipa columbiana See needlegrass, subalpine Tetradymia canescens See horsebrush, gray Stipa comata See needle-and-thread Tetradymia glabrata See horsebrush, littleleaf Stipa lettermanii See needlegrass, Letterman Tetradymia nuttallii See horsebrush, Nuttall Stipa occidentalis See needlegrass, western Tetradymia spinosa See horsebrush, spiny Stipa thurberiana See needlegrass, Thurber thimble seeder 79Ð80 Stipa viridula See needlegrass, green thimbleberry 266Ð267, 591Ð592, 601, 605 stork’s bill See alfileria Thinopyrum intermedium See wheatgrass, intermedi- Suaeda torreyana See desert blite; sumpbush, desert ate subalpine communities 601 Thinopyrum ponticum 320 See also wheatgrass, tall subalpine herblands 9, 201Ð202, 204Ð205, 259 threeawn complex 349 succession, plant 114 threeawn, fendler 310 successional changes 131 threeawn, purple 309Ð310, 349 suckers 762 threeawn, red See threeawn, purple sulfate-sulphur 52 timothy 132, 135, 142, 196, 205Ð206, 208, 215Ð216, sulphur 51, 54 259, 261, 264Ð265, 269, 271, 289, 309, 311, sumac, Rocky Mountain smooth 272, 276, 278, 283, 399Ð401, 701 604, 611Ð613, 711, 741, 752 Clair 401 sumac, skunkbush 135, 272, 276, 278, 283, 288, 604, Climax 302, 400 608Ð611, 711, 741, 752 Drummond 302 Bighorn 611 varieties 401 sumac, smooth 143 timothy, alpine 259, 269, 289, 309, 311, 398Ð399, 701, summercypress, Belvedere 262 763 sumpbush, desert 469, 491 timothy, mountain See timothy, alpine sunflower 287 tissue analysis 53 sunflower, annual 701 topography 9, 10, 11, 12, 13 surface seeder 80 toxicant 97 sweetanise 196, 216, 260Ð261, 270, 289, 426, 428, transplanter 82, 83, 740, 742 454, 704, 724, 728, 731, 741 transplanting 23, 262, 739 sweetclover, white 429 equipment 764, 765 sweetclover, yellow 135, 138, 196, 262, 264, 272Ð273, treatment of propagative material 190 275, 278Ð280, 283Ð284, 286, 429, 451Ð452, 704, treatments 724, 728 costs 64 Goldtop 452 economic benefits 64 Madrid 452 impacts on resources 64 Yukon 452 visual impacts 64 sweetroot, spreading 270, 429 trefoil, birdsfoot 272, 283 sweetvetch, northern See sweetvetch, Utah trencher 84

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-17 Trifolium See clover weeds, annual communities 308, 601 Trifolium fragiferum See clover, strawberry weeds, annual lowland communities 9, 257Ð258 Trifolium hybridum See clover, alsike weeds, noxious 211 Trifolium pratense See clover, red wet and semiwet meadow communities 9, 168, Trifolium repens See clover, white 206Ð208, 261, 308, 601 trisetum, spike 309, 311 wet meadows 262 Truax drill 74, 75 wheatgrass, beardless 310, 701 See also wheatgrass, Truax Seed Slinger 76 bluebunch Truax single row seeder 81 Whitmar 302 turkey 178, 179 wheatgrass, bluebunch 132, 135, 139, 142, 182Ð184, Tye drill 75 196, 271, 273, 275, 277, 280, 282, 284, 286Ð287, 294, 309Ð310, 337Ð342, 724, 728, 763 U Anatone 302, 341 U.S. Department of Agriculture (USDA) Goldar 341 Agricultural Research Service 4 Goldlar 302 Bureau of Plant Industry 2, 3 NewHy 302, 341 Division of Botany 2 P-7 302, 341 Forest Reserves 2 Whitmar 342 Forest Service 17 wheatgrass, bluebunch bearded 310, 701 Great Basin Experimental Range 16 wheatgrass, crested 182, 312, 316 See wheatgrass, Great Basin Station 3Ð4, 16, 17 fairway crested Intermountain Forest and Range Experiment wheatgrass, desert 182Ð184, 196 See wheatgrass, Station 3, 16 standard crested Intermountain Research Station 4Ð5, 17 wheatgrass, fairway crested 132, 135, 138, 142, Office of Grazing Studies 3 183Ð184, 196, 262, 271, 273, 275, 277, 279Ð280, Rocky Mountain Research Station 5 282, 284Ð287, 293, 294, 297, 297Ð318, 309, 310, Shrubland Biology and Restoration Project 4 315, 316, 701, 728 Natural Resources Conservation Service 5, 17 Douglas 302, 316 Plant Materials Centers 4 Ephraim 302, 316Ð317, 724 urea 54 Fairway 302, 317 Utah Division of Wildlife Resources 4, 17 Nordan 316 Utah State Division of Fish and Game 16 Parkway 302, 317 Ruff 302, 317 V wheatgrass, hybrid CD-II 316 vacuum-type seed harvesters 713 Hycrest 316Ð317 valerian, edible 260Ð261, 264, 429 NewHy 262 vegetation condition 26 SL-1 342 vegetation status 61 SL1 302 vegetation types 7, 9, 11, 13, 34, 36Ð38, 308, 601 See wheatgrass, hybrid crested also communities, plant CDII 303 vegetative propagation 748, 759Ð763 Hycrest 294, 303 vetch, American 182Ð183, 260, 270, 429 Kirk 303 vetch, bramble 429 wheatgrass, intermediate 132, 135, 138, 142, 182, Vicia americana See vetch, American 196, 221, 237, 255Ð256, 259, 269, 271, 273, 275, Viguiera miltiflora nevadensis See goldeneye, Nevada 277, 279, 280, 282, 284, 286, 287, 293Ð294, Viguiera multiflora multiflora See goldeneye, showy 309Ð310, 327Ð331, 701, 724, 727Ð728 violet, Nuttall 260 Amur 303, 330 virginsbower, western 605, 654Ð656, 711, 752 Chief 303, 330 vitamin A 179 Clarke 303, 330 W Greenar 330 Greenbar 303 water 171Ð172 Greenleaf 304, 331 water developments 171 Luna 304, 331, 724 watershed 131 Oahe 303, 330 weed control 62Ð63, 131, 147 Ree 330

Index-18 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Reliant 303, 330 wheatgreass, crested 297 Rush 303, 330 whortleberry 711 Slate 303, 330 wilding stock 739Ð740, 742, 759 Tegmar 303, 331 wildlife habitat 109Ð110, 112, 131, 155, 539 Topar 304, 331 cover 110, 131, 157 wheatgrass, pubescent 309 See also wheatgrass, edges, vegetative 111 intermediate security 162 wheatgrass, Siberian crested 142, 196, 246, 273, 275, thermal 161 279Ð280, 284, 286Ð287, 291Ð293, 309Ð310, 314, improvement 156 332Ð334, 701 management 113 P27 304 wildrye, alkali 309, 311, 382 See also wildrye, low Vavilov 304 creeping wheatgrass, slender 142, 196, 259, 264Ð265, 269, wildrye, Altai 309Ð310 271, 277, 282, 287, 289, 309Ð310, 342, 701 Eejay 306 Adanac 304, 344 Pearl 306 Primar 304, 344 Prairieland 306 Pryor 304, 345 wildrye, beardless 311 See wildrye, creeping Revenue 304, 345 wildrye, blue 269, 309Ð310, 377Ð379 San Luis 304, 345 Arlington 306, 378Ð379 wheatgrass, Snake River 309, 338 wildrye, Canada 309Ð310, 373Ð374 Secar 304, 338, 341, 342 Mandan 306, 374 wheatgrass, standard crested 132, 135, 138, 142, 271, wildrye, creeping 264, 265, 311, 383Ð384 273, 275, 277, 279, 280, 282, 284Ð287, 291Ð294, Rio 384 309Ð310, 313, 320Ð324, 724 Shoshone 384 Nordan 304, 324 wildrye, Dahurian Summit 304, 326 Arthur 306 wheatgrass, streambank 209, 220, 228, 262, 271, 273, James 306 275, 277, 279Ð280, 282, 284Ð287, 291Ð292, 294, wildrye, Great Basin 132, 135, 138, 142, 196, 246, 310, 320, 701, 763 See also wheatgrass, 261, 264Ð265, 275, 277, 279, 280, 282, 284, 287, thickspike 292, 309Ð310, 374Ð377, 702, 728 wheatgrass, tall 132, 135, 142, 196, 255, 261, 262, Magnar 306, 376Ð377, 724 264, 265, 271, 280, 292, 309, 310, 325Ð327, 702, Trailhead 306, 377 724, 731 wildrye, low creeping See wildrye, alkali Alkar 305, 326 wildrye, mammoth 264, 265, 309Ð310, 702 Jose 305, 327 Volga 306 Largo 305, 327 wildrye, purple 310 Orbit 305, 327 wildrye, Russian 132, 135, 138, 142, 196, 262, 264, Platte 305, 327 273, 275, 279Ð280, 284Ð287, 291Ð293, 309, 310, wheatgrass, thickspike 228, 237, 271, 275, 277, 279, 379Ð380, 702, 724, 728 280, 282, 284, 286Ð287, 294, 305, 309Ð310, Bozoisky-select 307, 380 318Ð320, 702, 763 Cabree 307, 380 Bannock 305, 319 Mankota 307, 380 Critana 305 Mayak 307, 381 Schwendimar 305, 320 Swift 307, 381 Sodar 305 Tetracan 307, 381 Soday 320 Vinall 307, 381 wheatgrass, western 135, 139, 142, 182Ð184, 220, wildrye, Salina 262, 291, 309Ð310, 381Ð382 228, 242, 255, 264Ð265, 271, 273, 275, 277, 280, wildrye, yellow 310 282, 284Ð285, 287, 291Ð294, 309Ð310, 334Ð337, willow 206, 208, 210Ð11, 213, 262, 266Ð268, 668Ð670 701, 763 willow, arroyo 268, 680Ð681 Arriba 305, 336 Rogue 681 Barton 305, 336 willow, barrenground 268 Flintlock 305, 336 willow, Bebb 268, 670Ð672 Rodan 305, 336 Wilson 672 Rosana 305, 336, 337 willow, Booth 268, 605, 672Ð673 Walsh 305, 337 willow, coyote 605, 676, 752

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-19 willow, Drummond 268, 605, 673Ð675 Hatch 484, 488 Curlew 674 winterfat, foothills 486 willow, Geyer 268, 676Ð677 winterfat, Pamirian 469, 486, 488 willow, grayleaf 268, 677Ð678 wipe-on applicators 92, 93 willow, Pacific 268 wormwood 495 See sage, Louisiana willow, peachleaf 268 wormwood, common 498 willow, plainleaf 683Ð684 wormwood, dwarf 498 willow, purpleosier 262, 605, 752 wormwood, oldman 495, 497Ð498, 605, 711, 741, 753 willow, sandbar 268 See also willow, coyote wortleberry, big 753 willow, Scouler 268, 605, 684Ð686, 752 willow, whiplash 605, 679Ð680 Y Nehalem 680 yarrow, western 260Ð261, 264, 270, 272Ð273, 275, Roland 680 279Ð280, 283, 287, 289, 291, 294, 426, 429, 430, willow, Wolf 268, 686Ð687 704, 741, 763 willow, yellow 681Ð683 yellowbrush See rabbitbrush winterfat 132, 135, 138, 142Ð143, 182Ð183, 185, 196, 201, 209, 228, 236Ð237, 242, 245Ð247, 250Ð251, Z 253, 274, 276, 278Ð279, 281, 284Ð285, 291Ð293, 469, 470, 472, 484Ð486, 605, 711, 725, 727, 730, zinc 51Ð52, 54 741, 752 zuckia, Arizona 469 winterfat communities 251 Zuckia brandegei See hopsage, spineless

Index-20 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 RMRS ROCKY MOUNTAIN RESEARCH STATION

The Rocky Mountain Research Station develops scientific information and technology to improve management, protection, and use of the forests and rangelands. Research is designed to meet the needs of National Forest managers, Federal and State agencies, public and private organizations, academic institutions, industry, and individuals. Studies accelerate solutions to problems involving ecosystems, range, forests, water, recreation, fire, resource inventory, land reclamation, community sustainability, forest engineering technology, multiple use economics, wildlife and fish habitat, and forest insects and diseases. Studies are conducted cooperatively, and applications may be found worldwide.

Research Locations

Flagstaff, Arizona Reno, Nevada Fort Collins, Colorado* Albuquerque, New Mexico Boise, Idaho Rapid City, South Dakota Moscow, Idaho Logan, Utah Bozeman, Montana Ogden, Utah Missoula, Montana Provo, Utah Lincoln, Nebraska Laramie, Wyoming

*Station Headquarters, Natural Resources Research Center, 2150 Centre Avenue, Building A, Fort Collins, CO 80526

The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the basis of race, color, national origin, sex, religion, age, disability, political beliefs, sexual orientation, or marital or family status. (Not all prohibited bases apply to all programs.) Persons with disabilities who require alternative means for communication of program information (Braille, large print, audiotape, etc.) should contact USDA’s TARGET Center at (202) 720-2600 (voice and TDD). To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, Room 326-W, Whitten Building, 1400 Independence Avenue, SW, Washington, DC 20250-9410 or call (202) 720-5964 (voice or TDD). USDA is an equal opportunity provider and employer.

Federal Recycling Program Printed on Recycled Paper Winterfat seedstalk Photo by Nancy Shaw